ENDODONTOE) LAND SNAILS FROM PACIFIC
ISLANDS
(Mollusca: Pulmonata: Sigmurethra)
PartH
Families Punctidae and Charopidae, Zoogeography
Alan Solem
Field Museum of Natural History
Chicago, Illinois
.
UNlVLhSiTY OF
ILLINOIS LIBRARY
AT. URBANA -CHAMPAIGN
ENDODONTOID LAND SNAILS FROM PACIFIC
ISLANDS
(Mollusca: Pulmonata: Sigmurethra)
Part II
Families Punctidae and Charopidae, Zoogeography
Alan Solern
Curator of Invertebrates
Field Museum of Natural History
Field Museum of Natural History
Chicago, Illinois
Submitted December 29, 1978
Published by:
Field Museum of Natural History
December 30, 1982
Mailed January 7, 1983
Library of Congress Catalog Card No.: 76-9516
ISSN 0015-0754
PRINTED IN THE UNITED STATES OF AMKRICA
II
CONTENTS
LIST OF FIGURES V
LIST OF TABLES VIII
INTRODUCTION 1
Acknowledgements 2
PREVIOUS STUDIES 3
MATERIAL STUDIED 4
METHODS OF ANALYSIS 5
PATTERNS OF MORPHOLOGICAL VARIATION 8
Size and shape variations 8
Shell sculpture 9
Apertural barriers 15
Parietal barriers 19
Columellar barriers 21
Palatal barriers 21
Patterns of barrier variation 23
Summary of barrier variations 23
Summary of shell variations 23
Gross anatomy 23
Genital system 24
Ovotestis 24
Hermaphroditic duct 24
Talon and carrefour 25
Albumen gland 25
Prostate and uterus 25
Terminal male genitalia 25
Terminal female genitalia 30
Pallial complex 30
Digestive system 32
Free muscle system 34
Nervous system 34
External body features 34
Patterns of visceral hump reduction 36
Summary of anatomical variation 37
CHAROPID-ENDODONTID CONCHOLOGICAL COMPARISONS 38
HABITAT RANGE AND EXTINCTION 45
PHYLOGENY AND CLASSIFICATION 46
FAMILY CLASSIFICATION OF THE ENDODONTOIDS 47
PHYLOGENETIC PROCEDURES 48
GENERIC CLASSIFICATION 49
SYSTEMATIC REVIEW 50
List of the taxa 50
Geographic keys to the genera 51
Family Punctidae 53
Genus Punctum Morse, 1864 57
Family Charopidae 59
Subfamily Rotadiscinae 70
Genus Microcharopa, new genus 70
Subfamily Charopinae 72
Genus Discocharopa Iredale, 1913 74
Genus Sinployea, new genus 81
III
IV
Society Island Sinployea 85
Cook Island Sinployea 94
Samoan and Swains Island Sinployea 117
Western Polynesian and Micronesian Sinployea 133
Fijian Sinployea 145
Melanesian Sinployea 166
Genus Ba, new genus 172
Genus Maafu, new genus 174
Genus Lauopa, new genus 177
Genus Tuimalila, new genus 178
Genus Lagivala, new genus 183
Genus Vatusila, new genus 191
Genus Graeffedon, new genus 200
Subfamily Trukcharopinae, new subfamily 205
Genus Trukcharopa, new genus 207
Genus Kubaryiellus, new genus 210
Genus Russatus, new genus 214
Genus Roimontis, new genus 217
Genus Palikirus, new genus 219
Genus Jokajdon, new genus 222
Genus Palline, new genus 228
Subfamily Semperdoninae, new subfamily 235
Genus Semperdon, new genus 236
Genus Ladronellum, new genus 255
Genus Himeroconcha, new genus 259
Incertae sedis 266
ZOOGEOGRAPHY 268
Patterns of family and generic distributions 268
Patterns of diversity 275
Hawaiian Islands 275
Marquesas Islands 277
Tuamotu Archipelago 279
Mangareva, Gambier Islands 280
Rapa Island 282
Austral Islands 287
Society Islands 288
Cook Islands 289
Samoa 290
Tonga 292
Lau Archipelago 292
Fiji, Main Islands 295
Ellice Islands 295
Marshall Islands 295
Caroline Islands 297
Mariana Islands 297
Palau Islands 298
Summary of specific and generic diversity 301
Shell size and geography 305
Hypothesized origin and radiation of Pacific Island endodontoid snail fauna 307
Patterns of species diversity 310
Summary of zoogeography 315
SUMMARY 316
REFERENCES 317
APPENDIX: Anatomical terms 324
INDICES . 325
LIST OF FIGURES
1. Shell microsculpture inSinployea modicella 10
2. Shell microsculpture inSinployea peasei and Tuimalila pilsbryi 11
3. Shell microsculpture inPalline biakensis 12
4. Shell microsculpture in Microcharopa mimula 13
5. Details of shell sculpture inDiscocharopa aperta 16
6. Microstructure of largest palatal barrier inJokajdon callizonus 18
7. Surface of largest palatal barrier in Graeffedon graeffei 19
8. Structure of apertural barriers in Semperdon xyleborus 20
9. Anatomy ofCharopa coma 26
10. Interior of lower vas deferens and epiphallus in Charopa coma 27
11. Anatomy of Phenacohelix pilula 28
12. Radular teeth ofPunctum minutissimum 33
13. Radular teeth of Tuimalila pilsbryi 35
14. Radular teeth of Tuimalila pilsbryi 36
15. Mean shell height distribution in the Endodontidae and Charopidae 38
16. Mean shell diameter distribution in the Endodontidae and Charopidae 39
17. Mean height/diameter ratio distribution in the Endodontidae and Charopidae 39
18. Mean whorl count distribution in the Endodontidae and Charopidae 39
19. Mean diameter/umbilical width ratio distribution in the Endodontidae and Charopidae 40
20. Mean spire height/body whorl width ratio distribution in the Endodontidae and Charopidae 40
21. Mean ribs on body whorl distribution in the Endodontidae and Charopidae 40
22. Mean ribs/mm, on the body whorl distribution in the Endodontidae and Charopidae 41
23. Patterns of whorl width increment in selected Endodontidae and Charopidae 42
24. Partial anatomy ofPunctum minutissimum 54
25. Anatomy ofLaoma leimonias 55
26. Partial anatomy ofParalaoma aprica 57
27. Anatomy of "Paralaoma" coesa 58
28. Punctum polynesicum 60
29. Anatomy of Flammulina zebra 64
30. Anatomy ofAmphidoxa marmorella 65
31. Anatomy ofStephanoda binneyana 67
32. Anatomy of Pseudocharopa lidgbirdi 69
33. Microcharopa mimula 71
34. Anatomy of Discocharopa aperta 75
35. Parietal barrier variation inDiscocharopa aperta 76
36. Discocharopa aperta from the New Hebrides 80
37. Australian Discocharopa aperta 81
38. Sinployea modicella and S. tahitiensis 87
39. Anatomy of Tahitian and Moorean Sinployea 90
40. Sinployea lamellicosta 91
41. Sinployea montana andS. neglecta 93
42. Anatomy of Society and Cook Island Sinployea 96
43. Anatomy of Sinployea avanaensis 97
44. Sinployea andrewi andS. atiensis 100
45. Sinployea peasei and S. avanaensis 103
46. Sinployea proximo and S. planospira 107
47. Sinployea decorticata andS. rudis 109
48. Sinployea harveyensis and S. youngi 113
49. Sinployea canalis 115
50. Sinployea otareae and S. tenuicostata 116
V
VI
51. Sinployea clista andS. clausa 120
52. Sinployea aunuuana andS. intermedia 123
53. Anatomy of Sinployea aunuuana, S. clista, and S. intermedia 124
54. Sinployea allecta allecta and S. a. tauensis 126
55. Anatomy of Sinployea allecta allecta 127
56. Sinployea complementaria 130
57. Anatomy of the Samoan Sinployea complementaria 132
58. Sinployea vicaria vicaria and S. v. paucicosta 136
59. Shell sculpture details of Sinployea vicaria vicaria 137
60. Anatomy of the Tongan Sinployea vicaria vicaria 138
61. Sinployea pseudovicaria andS. rotumana 139
62. Sinployea ellicensis ellicensis and S. e. nukulaelaeana 141
63. Sinployea kusaieana and Trukcharopa trukana 144
64. Sinployea recursa andS. angularis 148
65. Sinployea princei andS. irregularis 151
66. Sinployea inermis inermis, S. i. meridionalis, and S. i. lakembana 156
67. Anatomy of Fijian, Caroline Island, and New Hebridean Sinployea 158
68. Sinployea lauensis andS. navutuensis 160
69. Sinployea godeffroyana andS. adposita 162
70. Sinployea euryomphala 168
71. Sinployea solomonensis 169
72. Sinployea kuntzi 170
73. Sinployea novopommerana and S. descendens 171
74. Ba humbugi 173
75. Anatomy ofBa humbugi 175
76. Maafu thaumasius 176
77. Lauopa mbalavuana 178
78. Tuimalila pilsbryi and T. infundibulus 180
79. Anatomy of the Tongan Tuimalila pilsbryi 182
80. Lagivala vivus andL. minusculus 186
81. Lagivala microglyphis andL. macroglyphis 188
82. Vatusila kondoi and V. nayauana 193
83. Vatusila tongensis and V. niueana 197
84. Partial anatomy of Vatusila tongensis 198
85. Vatusila uaitupuensis and V. eniwetokensis 199
86. Graeffedon graeffei 201
87. Anatomy of the Samoan Graeffedon graeffei 203
88. Graeffedon savaiiensis and G. pricei 204
89. Kubaryiellus kubaryi andRussatus nigrescens 213
90. Anatomy ofRussatus, Kubaryiellus, and Trukcharopa 216
91. Roimontis tolotomensis andPalikirus cosmetus 218
92. Jokajdon callizonus and J. tumidulus 224
93. Anatomy of Jokajdon andPalline 227
94. Palline notera notera and P. n. palauana 230
95. Palline notera gianda andP. micramyla 233
96. Palline biakensis 235
97. Semperdon uncatus 239
98. Semperdon kororensis andS. xyleborus 242
99. Semperdon heptaptychius andS. rotanus 246
100 Anatomy of Semperdon 249
101. Ladronellum mariannarum 256
102. Anatomy of Ladronellum mariannarum 258
103. Anatomy ofHimeroconcha 261
104. Himeroconcha rotula and//, lamlanensis 262
105. Himeroconcha quadrasi and//, fusca 264
106. Distribution of Pacific Island endodontoid families 269
107. Distribution of most generalized and most specialized endodontid genera 270
108. Ranges of Lagivala and Vatusila 271
109. Range of Mautodontha and areas where umbilical brood chambers evolved 272
110. Range of Sinployea . 273
VII
111. Ranges ofGraeffedon, Palline, and Semperdon 274
112. Collecting localities on smaller islets of Mangareva, Gambier Islands 280
113. Collecting localities on Mangareva Islet, Mangareva, Gambier Islands 281
114. Collecting localities on Rapa Island 283
115. Major geographic features on Rurutu, Austral Islands 287
116. Collecting localities on Tubuai, Austral Islands 288
117. Collecting localities on Raivavae, Austral Islands 289
118. Collecting localities on Rarotonga, Cook Islands 290
119. Collecting localities on Upolu, Samoa 291
120. Collecting localities on Savai'i, Samoa 291
121. Main islands of Tonga 292
122. Collecting localities on Vava'u, Tonga 293
123. Collecting localities on Tongatapu, Tonga 294
124. Collecting localities on Eua, Tonga 295
125. Islands of the Lau Archipelago 296
126. Collecting localities on Kusaie, Caroline Islands 298
127. Collecting localities on Ponape, Caroline Islands 298
128. Islets of the Truk Group, Caroline Islands 298
129. Collecting localities on Moen Islet, Truk, Caroline Islands 298
130. Collecting localities on Tol Islet, Truk, Caroline Islands 299
131. Collecting localities on Dublon Islet, Truk, Caroline Islands 299
132. Collecting localities on Guam, Mariana Islands 299
133. Islands of the Palau Group 300
134. Collecting localities on Peleliu, Palau Group 300
135. Collecting localities on Koror and Auluptagel, Palau Group 300
136. Collection station on Angaur, Palau Group 301
137. Species diversity in island groups for the Endodontidae and Charopidae 303
138. Generic diversity in island groups for the Endodontidae and Charopidae 304
139. Median mean and mean ranges of adult shell diameter in island groups for the Endodontidae
and Charopidae 306
140. Species-area relationships for total land snail faunas of selected Pacific Islands 312
141. Species-area relationships for selected land snail families on Polynesian and Micronesian islands 313
142. Species diversity in islands grouped by size 313
143. Species diversity in islands grouped by elevation 313
LIST OF TABLES
I. Range of variation in the Pacific Island Charopidae 8
II. Body whorl contour in Endodontidae and Charopidae 8
III. Shell diameter and rib spacing in the Charopidae 14
IV. Degree of apertural narrowing by barriers 18
V. Percentage distribution of columellar barrier numbers 21
VI. Percentage distribution of palatal barrier numbers 22
VII. Palatal barrier numbers in constricted aperture Charopidae 22
VIII. Phyletic representation of dissected taxa 24
IX. Relative length of kidney lobes in the Pacific Island Charopidae 31
X. Conchological comparisons of Endodontidae and Charopidae from Pacific Islands 38
XI. Species numbers of Endodontoid land snails from selected geographic areas 53
XII. Range of variation in Punctum, Discocharopa, and Microcharopa 59
XIII. Geographic variation in Discocharopa aperta 79
XIV. Ratio of penis length to shell diameter in Sinployea 83
XV. Mean species measurements in Sinployea 83
XVI. Range of variation in Society Island Sinployea 85
XVII. Local variation in Society Island Sinployea, 1 89
XVIII. Local variation in Society Island Sinployea, II 95
XIX. Range of variation in Cook Island Sinployea 98
XX. Local variation in Cook Island Sinployea, 1 101
XXI. Local variation in Sinployea peasei 104
XXII. Local variation in Cook Island Sinployea, II Ill
XXIII. Local variation in Cook Island Sinployea, III 112
XXIV. Range of variation in Samoan and Swains Island Sinployea 118
XXV. Local variation in Samoan Sinployea, 1 121
XXVI. Local variation in Samoan Sinployea, II 128
XXVII. Range of variation in Western Polynesian and Micronesian Sinployea 133
XXVIII. Local variation in Sinployea vicaria 135
XXIX. Local variation in Western Polynesian Sinployea 142
XXX. Range of variation in Fijian Sinployea 146
XXXI. Local variation in Fijian, Melanesian, and Micronesian Sinployea 152
XXXII. D/U ratio frequency in Sinployea inermis subspecies 153
XXXIII. Ribs/mm, frequency in Sinployea inermis subspecies 153
XXXIV. Rib frequency in Sinployea inermis subspecies 153
XXXV. Local variation in Sinployea inermis, 1 154
XXXVI. Local variation in Sinployea inermis, II 155
XXXVII. Local variation in Sinployea adposita andS. irregularis 164
XXXVIII. Range of variation in Melanesian Sinployea 167
XXXIX. Range of variation in Fijian and Tongan endemic genera 181
XL. Range of variation inLagivala 184
XLI. Local variation inLagivala demani 191
XLII. Range of variation in Vatusila 194
XLIII. Range of variation in Graeffedon 202
XLIV. Sympatry of Charopidae on Ponape, Caroline Islands 208
XLV. Range of variation in Micronesian Charopinae, 1 209
XLVI. Local variation in Trukcharopa 211
XLVII. Age distribution in Trukcharopa trukana 211
XLVIII. Local variation in Kubaryiellus andRussatus 215
XLIX. Local variation inPalikirus andJokajdon 220
L. Range of variation in Micronesian Charopinae, II 223
VIII
IX
LI. Sculptural variation inJokajdon tumidulus 226
LII. Local variation inPalline 232
LIII. Range of variation in Semperdoninae 237
LIV. Local variation in Palau Island Semperdon 240
LV. Summary of barrier variation in Mariana Islands Semperdon 241
LVI. Local variation in Semperdon rotanus 248
LVII. Barrier variation in Semperdon rotanus 248
LVIII. Local variation in types and large form of Semperdon heptaptychius 252
LIX. Local variation in Semperdon heptaptychius 253
LX. Sculpture variation in Semperdon heptaptychius 254
LXI. Frequency distribution of diameter in populations of Semperdon heptaptychius 254
LXII. Frequency distribution of ribs in populations of Semperdon heptaptychius 254
LXIII. Frequency distribution of ribs/mm, in populations of Semperdon heptaptychius 254
LXIV. Barrier variation in Semperdon heptaptychius 255
LXV. Local variation in Ladronellum and Himeroconcha 257
LXVI. Generic distribution of Hawaiian Endodontidae and Punctidae 276
LXVII. Geographic distribution of species level taxa in Hawaiian Endodontidae 276
LXVIII. Diversity of Hawaiian land snails 276
LXIX. Land snail fauna of Rapa Island 284
LXX. Ages of Pacific Islands 307
LXXI. Pacific Island land snail families with comprehensive distributional data available 311
LXXII. Land snail faunas of Lord Howe, Upolu, and Viti Levu 312
LXXIII. Calculated and observed maximum density of land snail species 312
LXXIV. Correlation between island size, elevation, and number of land snail species 314
LXXV. Land snail diversity on islands of same size but different distances from Indonesian-Melane-
sian core 314
LXXVI. Diversity of land snails on small, isolated islands 314
INTRODUCTION
This is the second of two monographs revising the
endodontoid land snails of Polynesia, Micronesia, and
Fiji. Part I, covering the older and larger family En-
dodontidae, reviewed 185 species-level taxa (Solem,
1976b). Subsequently (Solem, 1977a) I described Pro-
toendodonta laddi and Cookeconcha antiquus from
Late Pleistocene strata in core drillings on Midway
Atoll and have three new subfossil Pseudolibera from
Makatea (Solem, in preparation B). Together with the
98 species-level taxa belonging to the Families Punc-
tidae and Charopidae diagnosed below, a total of 290
species-level taxa have been reviewed during this pro-
ject. Describing the extensive Hawaiian Island
monophyletic radiation of up to 290 unnamed, now
mostly extinct, endodontid taxa preserved in the Ber-
nice P. Bishop Museum is a task left to others.
The basic descriptive work essentially was com-
pleted in 1969, based on research supported by Na-
tional Science Foundation grants G-16419, GB-3384,
and GB-6779. This has been supplemented by modest
materials received subsequently. As a by-product of
the basic monographs, papers on a number of prob-
lematic groups (Solem, 1964a-b, 1968b, 1969b, 1972b,
1973c-d), extralimital endodontoids (Solem, 1958b,
1961, 1968c, 1970a, c, 1975, 1976a, 1977c, in prepara-
tion A), biogeography (Solem, 1958a, 1959a, 1968a,
1969c, 1973c, 1979a-b, 1981), endodontoid biology
(Solem, 1969a, d, 1974), and fossil land snails have
been prepared (Solem, 1977a, in preparation B; Solem
& Yochelson, 1979).
In the early 1970s, the new availability of scan-
ning electron microscopy (hereafter SEM) added a
major new dimension to this project. A series of reports
(Solem, 1970b, 1972a, c, 1973a-b, f, 1976a, 1977a-b)
explored its potential, and extensive additional data
has been incorporated into the major reviews. The first
monograph was submitted for publication in De-
cember, 1973, and was issued in due course (Solem,
1976b).
Finishing Part II has involved making very dif-
ficult decisions as to where and when to terminate
additional information seeking. The Endodontidae are
restricted to the Pacific Islands and thus this mono-
graph was easily circumscribed. Both the Punctidae
and Charopidae have their main centers of diversity
elsewhere, have been liberally supplied with generic
and family-level names based on shell features and
hunches, frequently have been rather poorly described
and often never illustrated (particularly Iredale, 1913,
1933, 1937a-c, 1939, 1941a-c, 1942, 1944, 1945), and
essentially are unknown anatomically except for the
data supplied by Hutton (1884a), Suter (see Refer-
ences), and Climo (1969a-b, 1970, 1971a-b, 1978) on
New Zealand taxa and contributions on South African
taxa summarized by Connolly (1939). I have dissected
and partially illustrated more than 75 extralimital
species in addition to those included below. This
attempt to work out extralimital relationships and to
assign subfamily and generic names is only a start
toward producing an overall phylogeny and classifica-
tion.
The limits finally adopted in Part II were: (1) to
include all extralimital species known to belong in
genera endemic to Polynesia, Micronesia, and Fiji; and
(2) to include anatomical data on extralimital taxa
only when needed to establish the subfamily units used
in the systematic review and biogeography. Thus, 10 of
the 98 taxa discussed here are found only in such areas
as the New Hebrides, Solomon Islands, Bismarck Ar-
chipelago, West Irian, and Indonesia (six species of
Sinployea, three species of Lagivala, and Palline
biakensis), whereas another species, Discocharopa
aperta (Mollendorff, 1888), occurs sporadically in the
area covered (Fiji, Samoa, Austral and Society Islands)
but has its main distribution from the Philippines to
Australia. Data are presented on type genera or species
of several nominate subfamilies from diverse areas
(New Zealand, Juan Fernandez), but no attempt is
made to provide an overall classification of the
Charopidae in this monograph. There still are too
many gaps in our knowledge of anatomy and shell
structure.
By the early 1970s there still remained a few prob-
lem taxa. Microcharopa and Discocharopa showed
many differences from the other Pacific Island taxa,
but I had not been able to determine their affinities
because only empty shells were available. In addition,
there was the problem of Stenopylis, a second Philip-
pine Islands to Australia taxon. Both Stenopylis and
Discocharopa were known to occur in Central Aus-
tralia. Planned fieldwork in Western and Central Aus-
tralia in early 1974 hopefully would yield live mate-
rial. Early dry season collecting near Alice Springs
produced only dead shells, but museum records sug-
gested that both genera had a wide distribution in
North Australia. The probability of subsequently ob-
taining live materials seemed excellent. In addition,
the possibility of field survey work in Micronesia for
the Office of Endangered Species arose, which could
have greatly expanded the anatomical coverage of the
SOLEM: ENDODONTOID LAND SNAILS
Trukcharopinae and Semperdoninae. This prospect did
not materialize, but in 1976 and 1977 additional field
and dissection work in Australia permitted allocating
Discocharopa, Stenopylis, and Microcharopa to family
units. The writing of introductory sections and the
biogeographic review, assigning table and plate num-
bers, and final editing were completed in 1978.
It is with a mixture of delight and regret that I
conclude this systematic project delight at the new
information and ideas produced, regret that the hand
of man has so altered the Pacific Island environment
that future work on most of these taxa and testing of
the new ideas presented has been rendered impossible
by extinctions. The charopids apparently have been
less affected than the Endodontidae, but even with
these it is probable that many already are a vanished
group of organisms.
Concluding this monograph is a detailed biogeo-
graphic review of the endodontoid land snails. This
project spanned the time from the dominance of fixed
continents as a biogeographic axiom to the current
dogma of wandering continents. Land snails have not
been versed in biogeographic controversies, and their
patterns of distribution continue to fit neither model
exclusively.
The endodontoid land snails were the most spe-
ciose land snail group on the Pacific Islands (Solem,
1976b, pp. 1-2). They still are the most speciose group
in Australia and New Zealand (table XI), areas in
which they seem to be flourishing despite human ac-
tivities. The data and ideas presented in these volumes
hopefully will inspire more investigations of these
taxa.
ACKNOWLEDGEMENTS
In addition to the many individuals listed by
Solem (1976b, pp. 5-6), the following people have con-
tributed significantly to the success of this project. For
access to specimens in collections, suggestions as to
collecting localities, loan of materials, and many help-
ful ideas, I am indebted to Dr. Frank Climo, National
Museum of Natural History, Wellington, New Zealand;
Drs. Edmund Gittenberger and A. C. van Bruggen,
Rijksmuseum van Natuurlijke Historie, Leiden; Dr.
Jackie van Goethem, Institut Royal des Sciences
Naturelles de Belgique, Brussels; Dr. Simon Tillier,
Museum National d'histoire Naturelle, Paris; Dr.
Peter Mordan, British Museum (Natural History),
London; Dr. Barry Wilson, Dr. Fred Wells, Ms. Shirley
Slack-Smith, Mr. George Kendrick, Western Austra-
lian Museum, Perth; Dr. Winston Ponder, Australian
Museum, Sydney; and Mr. and Mrs. Fred Aslin, Mt.
Gambier, South Australia.
For assistance in the field, or supplementary
fieldwork by, I continue to be deeply in debt to Mr.
Laurie Price, Kaitaia, New Zealand, Field Associate of
Field Museum of Natural History. Mr. Carl Christen-
sen, University of Arizona; Dr. Barry Wilson, Shirley
Slack-Smith, George Kendrick, Clayton Bryce, and
many other staff members of the Western Australian
Museum; Fred and Jan Aslin, Mt. Gambier, South
Australia; Dr. Brian J. Smith, National Museum of
Victoria, Melbourne; Dr. Winston Ponder, Australian
Museum, Sydney; and Mr. Ron Kershaw, Launceston,
Tasmania, have assisted in Australian fieldwork or
have provided critical help with collecting localities.
Charts and figures in the introductory sections
have been prepared new or reworked by Ms. Elizabeth
Liebman, Illustrator, Division of Invertebrates, Field
Museum of Natural History. Invaluable help with
mounting and labeling was provided by Mrs. Dorothy
Karall, Associate, Division of Invertebrates. Illus-
trations for the systematic review were done primarily
by the several artists cited by Solem (1976b, p. 6). The
credit for the individual figures is indicated by the fol-
lowing abbreviations: Ms. Nelva Bonucchi (NB); Mrs.
Jane Calvin (JC); Mrs. Carole W. Christman (CW); Mr.
Sam Grove (SG); Mrs. Lynda Hanke (LH); Mr. Sander
Heilig (SH); Dr. Harold J. Walter (HJW); Mr. E. John
Pfiffner (EJP); Mrs. Patricia Rill Smiley (PS); Ms.
Marcia Oddi (MO); and the late Mrs. Margaret Anne
Moran McKibben (MM). In particular, the work by the
late Margaret Anne Moran McKibben was critical to
the success of this project.
Tabular material in the systematic review was
typed by Mrs. Sandra Rendleman and Ms. Jayne
Freshour. Tabular material in the introductory sec-
tions was typed by Mrs. Valerie Connor-Jackson, Sec-
retary, Division of Invertebrates.
Grateful acknowledgement is made to the Na-
tional Science Foundation, whose generous support
through grants G- 16419, GB-3384, GB-6779, and DEB
75-20113 in support of fieldwork, illustration prepara-
tion, and museum study of types has made this study
possible. The establishment of a scanning electron
microscope laboratory at Field Museum of Natural
History was funded by National Science Foundation
grant BMS 72-02149. Publication of Part I (Solem,
1976b) was in large part made possible by National
Science Foundation grant DEB 75-14048.
Much help with final editing and complete typing
of the manuscript was given by Ms. Sharon Bacoyanis
and Mrs. Valerie Connor-Jackson. Their efficient in-
volvement in every phase of final preparation was crit-
ical. Proofreading and indexing was greatly aided by
Valerie Connor-Jackson.
As with Part I, the years of labor by the late C.
Montague Cooke, and by Yoshio Kondo, plus the
facilities provided by the Bernice P. Bishop Museum
during 1961-1962 and their long loan of needed mate-
rials were instrumental to the initiation and comple-
tion of this project. Dr. Cooke had not reviewed any of
the taxa covered in this report, so all errors are my
responsibility.
PREVIOUS STUDIES
A more extensive historical review regarding the
Pacific Island taxa was given in Solem (1976b, pp. 7-8)
and should be consulted for background comments.
Previously described charopid taxa date from the
studies of Ferussac (1840), Hombron & Jacquinot
(1841), Mousson (1865, 1869, 1870, 1871), Cox (1870),
Pease (1870), Garrett (1872, 1881, 1884, 1887a-c),
Liardet (1876), Tapparone-Canefri (1883), Mollendorff
(1888, 1900), Beddome (1889), Quadras & Mollendorff
(1894), Smith (1897), Clapp (1923), Rensch (1937), Dell
(1955), Ladd (1958, 1968), and Solem (1959a, 1960). Of
the 45 previously described species-level taxa, 11 were
named by Garrett, six by Mollendorff, five by Mousson,
four by Quadras & Mollendorff, and four extralimital
species by Rensch. The remaining 15 species are scat-
tered among the other authors. The dates of species
descriptions were summarized in Solem (1976b, p. 3,
table III). The cluster of 1890 and 1900s taxa in the
Charopidae reflects the activities of Mollendorff and
his field associate, J. F. Quadras. The 10 taxa described
since 1920 include the highly significant fossils from
Eniwetok, Vatusila eniwetokensis (Ladd, 1958), and
Funafuti, Lagivala davidi (Ladd, 1968), plus basically
extralimital taxa (Solomon Islands, New Hebrides,
Bismarck Archipelago) that happen to belong to gen-
era also represented on the Pacific Islands. As with the
Endodontidae, the greatest period of descriptive activ-
ity was in the 1870s and 1880s, reflecting the publica-
tions of Mousson and Garrett.
No data had been recorded in the literature con-
cerning the anatomy of the Pacific Island taxa except
for a few notes on the tail, jaw, and radula ofRussatus
nigrescens (Mollendorff, 1900, p. 108, figs, 1-3) and the
radula and penis complex exterior of Sinployea
euryomphala (Solem, 1959a, pi. 6, figs. 7-8).
Because the main center of the charopid radiation
is extralimital to the Pacific Islands, extensive refer-
ence has had to be made to taxa from other parts of the
world. New Zealand, Australian, and New Caledonian
taxa in particular have yielded important comparative
data. Rather than detail the many reports here, cita-
tions are given under the organs or species referred to
at that place in the text. Here it is necessary only to
point out that these are diverse faunas with a long
record of study. The literature is highly fragmentary
and widely scattered, but fortunately there are sum-
mary references that enable relatively quick access to
the literature. For New Zealand, the classic mono-
graph of Suter (1913) and the checklist of Powell (1976)
summarize the work done prior to the highly signifi-
cant studies of Climo (1969a-b, 1970, 1971a-b, 1978).
For Australia, the checklist of Iredale (1937a-c) and
his faunistic reviews of nearby areas as well as Aus-
tralia (Iredale, 1939, 1941a-c, 1942, 1944, 1945) pro-
vide essential but very difficult to use reports. Franc
(1957) and Solem (1961) supplement the excellent re-
view of New Caledonian land mollusks given by Crosse
(1894).
Early observations on the anatomy and radular
structure of some Australian (Hedley, 1891, 1893) and
New Zealand (Hutton, 1884a, Suter, 1890, 1891a-b,
1892a-c, 1893a-d, 1894a-d, 1901, 1903, 1913)
charopids were accurate within the limits of optical
observation and the state of knowledge concerning
molluscan anatomy extant at that time, but have little
value today in deciding generic and family limits.
Even more than for the Endodontidae, the classifi-
cation of the charopids used previously was based on
form "pigeon holes," and the assignments made to gen-
era were based upon superficial conchological simi-
larities. Of the 21 genera reviewed below, only two
were named previously, and both of them, Punctum
Morse, 1864, and Discocharopa Iredale, 1913, are gen-
era whose main distributions lie outside the Pacific
Island area. All the remaining species belong to unde-
scribed, endemic taxa, mostly not yet known from
other than the fringing islands and Pacific Island area
proper.
MATERIAL STUDIED
The 98 taxa formally described or alluded to as
probably representing new taxa were represented by
about 7,500 specimens. Since the summary presented
by Solem (1976b, p. 9), a number of charopids from the
Lau Archipelago of Fiji collected by Price in 1970 and
Cernohorsky in 1977, Society Islands by Solem in 1977,
Western Australia by Solem, L. Price, and Carl Chris-
tensen in 1976 and 1977, and near Alice Springs in the
Northern Territory in 1974 and 1977 have become
available. Data from them has been incorporated into
the systematic review. No attempt at recalculating the
data in Solem (1976b, pp. 9-10, tables IV- V) has been
made because the numbers added are small in com-
parison to the previously listed materials.
Throughout the text, the following abbreviations
are used to indicate the repository of the cited speci-
mens:
AIM Auckland Institute and Museum, Auckland
AMS Australian Museum, Sydney
ANSP Academy of Natural Sciences, Philadelphia
BMNH British Museum (Natural History), London
BPBM Bernice P. Bishop Museum, Honolulu
DMW National Museum of Natural History, Wellington
FMNH Field Museum of Natural History, Chicago
IRB Institut Royal des Sciences Naturelles de Belgique,
Brussels
JDCP Collection of Journal de Conchyliologie, Paris
MCZ Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge
MSNG Museo Civico di Storia Naturale "Giacomo Doria,"
Genova
NHM Naturhistorisches Museum, Basel
NMWC National Museum of Wales, Cardiff
RNHL Rijksmuseum van Natuurlijke Historic, Leiden
RSM Royal Scottish Museum, Edinburgh
SAM South Australian Museum, Adelaide
SMF Natur-Museum Senckenberg, Frankfurt-a.-M.
UMMZ University of Michigan, Museum of Zoology, Ann
Arbor
USNM National Museum of Natural History, Smithsonian
Institution, Washington, D.C.
WAM Western Australian Museum, Perth
ZMA Zoologisch Museum, Amsterdam
ZMB Zoologisches Museum der Humboldt-Universitat,
Berlin
Zurich Zoologisches Institut der Universitat, Zurich
Most of the material in the BPBM resulted from
the historical activities of Andrew Garrett plus speci-
mens obtained during three major expeditions in the
1930s. The Mangarevan Expedition from April 15
through October 28, 1934 (Cooke, 1935), Micronesian
Expedition from December 8, 1935, to June 10, 1936
(Gregory, 1936, p. 40), and the Henry G. Lapham Ex-
pedition to Fiji from June 27 through September 28,
1938 (Buck, 1939, pp. 29-30), contributed major mate-
rials for this study. Additional collections made in Fiji
by Harry G. Ladd, plus miscellaneous samples taken
by a variety of BPBM staff members, combined with
the above to provide the bulk of material analyzed.
Fieldwork sponsored by FMNH and National Sci-
ence Foundation in Rarotonga (1964 and 1965), West-
ern Samoa (1965), Fiji (1965 and 1970), Tonga (1965-
1966), New Caledonia (1967), and New Hebrides (1972)
by Laurie Price and/or Alan Solem was designed to
sample areas that were underrepresented in the BPBM
collections. Fieldwork in New Zealand (1962, 1965)
and various parts of Australia (1962, 1965, 1973, 1974,
1976-77) provided critical comparative materials that
enabled assigning subfamily names and indicating
possible phylogenetic trends within the Arionacea.
Only a few of these dissections are illustrated or cited
here because this report is deliberately restricted to the
Pacific Islands, except where a genus extends extralim-
itally.
Special features regarding the methods of collect-
ing and specimen storage used by the BPBM staff were
reviewed by Solem (1976b, p. 10) and are not repeated
here, although applying in full to the studied materials
used in this report.
METHODS OF ANALYSIS
The basic data on distribution of size and shape
variation within a population, how to recognize an
adult shell, the limits of measurement reliability, and
definitions of the standard measurements used were
fully explained in Solem (1976b, pp. 11-15).
It is necessary to emphasize some data concerning
sample bias because rather significant mean size
differences among museum lots of previously described
species are sometimes ignored. Prime examples of this
are in such Rarotongan taxa as Sinployea proxima
(Garrett, 1872) (table XX), S. rudis (Garrett, 1872)
(table XXIII), S. harveyensis (Garrett, 1872) (table
XXII), and the Funafuti Atoll S. ellicensis ellicensis
(table XXIX). The pattern of trading samples from
larger series to other museums or collectors still is
common practice among conchologists. Larger speci-
mens will tend to be retained by the original source,
with smaller examples being dispersed widely. The
possible extent of such dispersal is perhaps best illus-
trated by the endodontid land snail species Libera
tumuloides (Garrett, 1872). Although "over 300 speci-
mens" were collected originally, only 72 could be lo-
cated for this study, indicating dispersion and loss or at
least absence from major museum collections of more
than 228 examples in less than 100 years (Solem,
1976b, p. 430). In contrast, the endodontid Mauto-
dontha (M.)zebrina (Garrett, 1874) was known from "a
half dozen specimens," all of which were located
(Solem, 1976b, p. 162). Hence the reliability of mean
measurements made on rare species is apt to be much
greater than the means for common species where
trading bias probably has distorted the data base. The
exact extent of this distortion is unknown because the
original measurements were generally of only a single
specimen and were cited to the nearest millimeter. It
must be recognized that sizes cited here for pre-1900
species may be larger than the means of the popula-
tions from which they came, but how much larger is
unknown.
Data from the BPBM and FMNH collections in-
volved measuring field samples that had not been di-
luted by trading activities. Thus, these results are di-
rectly comparable with each other.
For each adult specimen, shell height, diameter,
whorl count, umbilical width, number of radial ribs on
the body whorl, number of apical spiral cords, and any
information concerning number and position of aper-
tural barriers were recorded. The only difference in
data recording from that used in Solem (1976b) is the
consistent counting of body whorl ribs and apical cords.
The lack of apertural barriers in many charopids
suggested greater use of sculptural features, and thus
body whorl ribs were counted wherever possible. Solem
(1976b, p. 14, fig. 5; p. 43, fig. 32; p. 52, fig. 36) defined
each of these parameters except for the apical spiral
cord count because the Endodontidae normally lacks
this type of major sculpture. This count was taken on a
suture-to-suture transect just before the end of the
apex, provided that portion of the shell was unworn. At
the start of the apex the exposed portion of the whorl
profile is narrower than at the end, producing a sig-
nificantly lower spiral ridge count. In Microcharopa
mimula (fig. 4a), for example, the apex tip shows 15
ribs, and the apex end, 20 ribs. Usually, if a count could
not be made near or at the usual position, the apex was
sufficiently worn so that no count could be obtained.
Apical cord count could not be tallied on many speci-
mens. It was, in general, subject only to minor error in
tallying where countable.
Height/Diameter (hereafter H/D) ratios, Diameter/
Umbilical Width (hereafter D/U) ratios, and Ribs/mm,
of body whorl circumference were calculated, the latter
by the formula:
rib count on body whorl
Ribs/mm. =
TT x shell diameter in mm.
The validity of this index was discussed by Solem
(1976b, pp. 42-43).
Means, ranges, and standard errors of the means
were computed for each parameter except whorl count
for each population. Some of these results are pre-
sented in tables headed "Local Variation in ... ."
The total variation within each species or subspecies is
summarized in tables headed "Range of Variation in
. . . ." Means are given, but no standard errors of the
mean were calculated because of the wide temporal
span (1820 to 1977) of the materials represented.
At a later stage in this review, from eight to 12
examples of each species, where sufficient materials
were available, were measured as to spire elevation
and body whorl width, with a suitable ratio (SP/BWW)
calculated and included in the "Range of Varia-
tion. . . ." tables. This gives an index of actual spire
elevation undistorted by differences in the degree of
body whorl descension behind the lip (see Solem,
1976b, pp. 24-26). At the same time these additional
measurements were made, the pattern of the micro-
sculpture on the early portion of the body whorl (see
Solem, 1976b, p. 42, fig. 32b) and the pattern of rib
spacing were determined and added to the draft diag-
noses and descriptions.
SOLEM: ENDODONTOID LAND SNAILS
Dissections of as many taxa as possible were car-
ried out between 1961 and 1977, with several rechecks
of structure as my understanding of the variation pat-
terns increased. Because only fragmentary preserved
animals were available for many species (see Solem,
1976b, p. 19), often only data on the terminal portions
of the genitalia could be presented. Where possible,
usually both the entire genitalia and the penis complex
interior plus the pallial region were illustrated. Data
on body color, tail structure, muscle system, and radular
structures were obtained, but have not been illustrated
because of lesser information content for classification
and phylogeny. Unfortunately, the bulk of the dissec-
tion, illustration, and descriptive work was done prior
to the availability of scanning electron microscopy as a
research tool. Hence only a modest number of SEM
observations are included in this study, although ex-
tensive use has been made of the SEM in working with
materials from extralimital areas.
The data base consists of many shell observations
and measurements, data from the pallial region,
genitalia, radula, and external body features. In the
previous report, some use was made of computer-
generated phylogenies (Solem, 1976b, pp. 114117,
figs. 59-61). No such attempts were made in studying
the Charopidae, because major features such as the
apertural barriers obviously arose independently in
several lineages (Solem, 1973b, p. 305), there were
sharp anatomical differences between major groups
with highly similar shell features, and the resulting
convergences would utterly confuse any phenetic pro-
gram unless de post facto elimination of convergences
was undertaken. The benefits of the computer model-
ing outlined by Solem (1976b, pp. 114-117) were
applied to this analysis without the work of program-
ming and running various combinations. Mean mea-
surement data was key-punched and numerous print-
outs prepared, but this was essentially an aid to the
phylogenetic analyses. Cladistic analysis was not at-
tempted, because there are still numerous gaps in
knowledge of the extralimital taxa. Outgroup compari-
sons would have been sketchy or incomplete at best,
since the large charopid and punctid faunas of New
Zealand, Australia, New Caledonia, and associated is-
lands occur under conditions of complex sympatry,
with many interspecific adjustments in penial chamber
features that make interpreting relationships exceed-
ingly difficult.
The result is that the proposed phylogeny has been
arrived at by phylogenetic methodology after looking
at a large number of characters in several organ sys-
tems and interpreting these according to the basic
three-tiered pattern of evolutionary events outlined in
Solem (1978b). I have used a pragmatic rather than an
ideologic approach.
The problem of how to determine species limits
when allopatric island populations are involved is
quite difficult (Solem, 1976b, pp. 15-17). Sympatric
congeneric species pairs or trios are invaluable in pre-
dicting the probability that two allopatric populations
are or are not reproductively compatible. Since the
Charopidae, as do the Endodontidae, use the surface
features of the penis chamber and lower female tract
for species recognition when two closely related species
are sympatric, there usually are demonstrable dif-
ferences in this region of the reproductive tract. Corre-
lated with this will be minor to prominent shifts in
conchological structures. By determining the "mini-
mum conchological difference" between sympatric con-
generic species pairs, a pragmatic standard is available
against which to measure the degree of conchological
difference between allopatric populations. If they equal
or exceed the "minimum conchological difference,"
they are presumed to have diverged genetically to the
level of species. If they show slight differences that are
below the "minimum conchological difference," they
are judged to be conspecific or at most subspecifically
separable. Under conditions of allopatry there are few
to no selective pressures for changes in the species rec-
ognition features of the genitalia. Indeed, there proba-
bly is rather strong selective pressure for maintaining
a conservative pattern of species recognition features
because departure from the pattern might be a mating
disadvantage.
This methodology does not presume genetic link-
age between conchological differences and species rec-
ognition features. Quite the contrary pattern is
hypothesized. The conchological differences are linked
to physiological or behavioral traits that have been
selected for by environmental pressures, producing
eventual basic genetic incompatibility. Upon becoming
microsympatric with a congener, strong selective pres-
sure for species recognition changes would exist, if, as
seems probable, nonmultiple mating is a standard pat-
tern for the charopids. Although we have no data on
the frequency of mating, the absence of any evidence of
more than one set of transferred sperm during dissec-
tion of several hundred individuals suggests that lim-
ited mating, rather than multiple mating, is more
probable.
There are far fewer clear examples of congeneric
sympatry with adequate materials for dissection in the
Charopidae than in the Endodontidae (see Solem,
1976b, pp. 80-83, tables LII-LIV). Partly this reflects
the difference in the distribution and probable age
within the area. Partly this reflects a greater vertical
range in habitat for the Charopidae, with tree trunk to
semiarboreal habitat available even at dryer eleva-
tions for the charopids as contrasted with the terres-
trially limited endodontids. Partly it is a matter of the
Charopidae, except for Sinployea, showing rather low
speciation but high generic diversity within the Pacific
Island area. The one island on which an apparent mul-
titude of species in one genus existed, Rarotonga, ap-
parently has nine of 11 charopid species extinct.
The example that comes closest to being sympatric
and showing minimal conchological differences in-
volves Sinployea peasei and S. avanaensis on Raro-
METHODS OF ANALYSIS
tonga. The shell of S. avanaensis (fig. 45d-f) has more
than twice the number of radial ribs on the body whorl,
a distinctly narrower umbilicus, and averages about
one-quarter whorl less, although it is nearly identical
in size and shape. Anatomically, the penis of S.
avanaensis is 1.41.6 mm. long, compared with 1.1-1.3
mm. in S. peasei; the verge (PV, fig. 43e) and muscle
collar of S. avanaensis are larger than the equivalent
structures in S. peasei (fig. 42d-e); and the pallial re-
gion of S. avanaensis (fig. 43b) is slightly shorter, with
a noticeably shorter and broader rectal kidney arm
than in S. peasei (fig. 42c). The two species apparently
have ecological separation, with S. peasei found almost
without exception under rotting wood and stones,
whereas, except at a very dry station, S. avanaensis
lived in moss and lichens on tree trunks. More detailed
collecting in Avana Valley on Rarotonga is needed to
determine if they are actually sympatric at any point
or if they are microallopatric in their distribution.
In the Lau Archipelago of Fiji, for example, both S.
inermis (Mousson, 1870) and S. adposita (Mousson,
1870) have been collected on Lakemba, Aiwa, and
Yangasa Levu, and fragmentary material adequate for
dissection of both species from a single island was
available. The penis length of S. inermis is 0.8-1.0
mm. compared with 1.65 mm. for S. adposita, and the
shells are very distinctive. The differences are larger
than in the Rarotongan example. Sinployea adposita
has a normal vergic papilla and a very small stimula-
tory pad, whereas S. inermis has a large stimulatory
pad and a small vergic papilla.
On Mt. Lamlan, Guam (Station 137), both Himer-
oconcha lamlanensis and H. fusca (Quadras & Mollen-
dorff, 1894) were taken alive. The former has a short-
ened penis (2.0 mm.) with major pilasters forming
halfway from epiphallic pore to atrium, and the latter
has a longer (2.7 mm.) penis with the major pilasters
starting just below the epiphallic pore (compare fig.
103a-b, d, f). These differences would serve quite effec-
tively as species recognition devices.
Fortunately, differences between species in the
Pacific Island Charopidae appear to be rather striking
and uniform, although often difficult to characterize
verbally because of extensive local variation.
Basically the same procedures and philosophical
approach were used to study both the Charopidae and
Endodontidae, so that the results are directly compar-
able in terms of species discrimination, classification,
and phylogenetic interpretations. The greater generic
diversity and usually lower specific diversity in the
Pacific Island Charopidae are interpreted as resulting
from multiple invasions from other areas at a more
recent time than for the Endodontidae, plus the proba-
ble fact that the Helicarionidae and Euconulidae may
have been invading only slightly later in time than the
Charopidae. Conceivably, this often arboreal group of
Limacacea could have denied the opportunity for ex-
tensive arboreal radiation to the Charopidae.
PATTERNS OF MORPHOLOGICAL VARIATION
Initial studies from 1961 through 1969 on both
shell and anatomy were carried out at 100 x magnifica-
tion or less using a Leitz dissecting microscope. Radu-
lar mounts in Euparol were examined unstained using
a Leitz Ortholux compound microscope under bright-
field, dark-field, and phase-contrast illuminations.
Subsequently, SEM observations on both shell and
radular features resulted in a series of reports (Solem,
1970b, 1972a-c, 1973a-b, d, 1974, 1975, 1976a,
1977a-b) whose results are partially incorporated here.
None of the available anatomical material was
suitable for histological investigation. In many cases
the apical genitalia of the BPBM material could not be
studied because of the preservation technique utilized
(Solem, 1976b, p. 19).
The following survey of shell and anatomical vari-
ation is less detailed than that presented for the En-
dodontidae (Solem, 1976b, pp. 19-99). Instead of one
phylogenetic lineage being represented, the Pacific Is-
land Charopidae include several independent lineages.
Instead of a monophyletic origin for the apertural bar-
riers, several independent origins are involved. Thus,
certain of the questions asked in the endodontid review
are not relevant and were omitted. In terms of shell
structure, the Charopidae on the Pacific Islands simply
are more conservative.
SIZE AND SHAPE VARIATIONS
At least one specimen was seen of all the species
treated in this monograph, so that the discussion that
follows includes all the named taxa. Specimens that
were indicated as probably representing new taxa, but
which were not formally named, have been omitted
from the comments and statistical analyses. No at-
tempt at use of factor or multivariate analysis has been
attempted because only the relatively crude standard
shell measurements were available. The overall pat-
tern of mean measurements of each species follows that
of the Endodontidae (Solem, 1976b, pp. 20-21, figs.
6-10) except that they are more centrally clustered.
Table I summarizes the basic distribution for nine var-
iables or ratios. Although the extremes are well re-
moved from the median, the distance between the first
and third quartiles is relatively restricted. For shell
height, diameter, and H/D ratio the first to third quar-
tile range is 18%- 19% of the total range, for the whorl
count it is only 13%, for apical cords only 16%, and for
the number of ribs on the body whorl (assuming the
maximum as 200) and body width it is 22.5%. Consid-
TABLE I. - RANGE OF VARIATION IN THE PACIFIC ISLAND CHAROPIDAE.
1st
Minimum Quartile
3rd
Quartile
Means of:
Height
0.
48
1.
23
1.
51
1
.81
3,
69
Diameter
1.
07
2.
27
2.
76
3
,49
7,
52
H/D Ratio
0.
365
0.
486
0.
523
.566
0.
801
Whorls
3
1/8+
3
7/8
4
1/8-
4
3/8-
6
5/8
D/U Ratio
2.
03
3.
44
3.
94
4.
73
CLOSED
Ribs on
body whorl
18
.9
74
.0
91
.1
115.0
REDUCED
Ribs /mm.
Body Whorl
Width
1.70
0.32
Apical Cords 6.5
7.66
0.76
9.5
11.25
0.90
10.5
14.61
1.15
12.0
REDUCED
2.06
22.0
ering that four phyletic lines are clumped, this is a
quite narrow basic range of variation. It is also much
less than in the Endodontidae (Solem, unpublished
data). The extent to which this results from distortions
of the brood chamber formation as opposed to the much
larger number of taxa and greater age of the Endodont-
idae is unknown.
Body whorl contour in the Charopidae shows the
same type of variation found in the Endodontidae but
differs in frequencies. Table II contrasts the two
families, using the same definitions as for the char-
acter states in the Endodontidae except for omitting
the brood chamber condition that has no Charopidae
equivalent. Lateral compression, angulation, or cari-
nation of the periphery is far more prevalent in the
TABLE II. - BODY WHORL CONTOUR IN ENDODONTIDAE AND CHAROPIDAE.
Total Number of Taxa:
Endodontidae
38
43
21
Body Whorl Contour
Laterally compressed
Evenly rounded
Flattened above and/or
below rounded periphery
Angled periphery 14
Carinated periphery 28
State not recorded for several taxa.
Charopidae
6
41
39
1
3
PATTERNS OF MORPHOLOGICAL VARIATION
Endodontidae. The Charopidae almost all have either
an evenly rounded periphery or a mild to prominent,
Sinployea canalis (Garrett, 1872) (fig. 49b), lateral flat-
tening above a rounded periphery. A very few species,
such as Sinployea planospira (Garrett, 1881) (fig. 46e),
have a clearly laterally compressed periphery. Only
Sinployea angularis (fig. 64e) has an angled periphery,
whereas Himeroconcha quadrasi (Mollendorff, 1894)
(fig. 105b), H. fusca (Quadras & Mollendorff, 1894) (fig.
105e), and especially Maafu thaumasius (fig. 76b) have
carinated peripheries. These differences in peripheral
contour obviously are not exclusive and have no value
in trying to characterize the families, except to point
out another area in which their average pattern of
growth diverges. In the Endodontidae there was a
noticeable effect on shell diameter and H/D ratio with
changes in body whorl contour (Solem, 1976b, p. 23, fig.
14). The same pattern applies to the few Charopidae
with drastically altered contours, but the numbers are
too few to warrant statistical treatment.
Because there are significant differences between
the families in terms of spire protrusion, a comparison
is given on pp. 38-40. The only really high-spired
charopid is Ba humbugi (fig. 74b). The only truly
sunken spire in the Charopidae is found in Roimontis
tolotomensis (fig. 91a), three additional species have
the spire depressed below the body whorl (Maafu
thaumasius, fig. 76a-b; Himeroconcha lamlanensis,
fig. 104d-e; Kubaryiellus kubaryi, fig. 89a-b), and 11
taxa have the spire barely emergent to slightly de-
pressed or actually variable in the case of Lagivala
demani (Tapparone-Canefri, 1883). The differences in
spire elevation are so minor that no analyses of possi-
ble effects on basic measurements are presented.
Umbilical shape in the Charopidae is monoton-
ously V-shaped, becoming saucer-shaped when more
widely open as in Discocharopa (fig. 36c) and nearly
U-shaped in the few species where it is narrowed (figs.
41c, 51f). There is none of the rich variation seen in the
Endodontidae (Solem, 1976b, p. 27, fig. 16), hence only
a few comments were presented in the family compari-
sons (p. 40).
Many of the Charopidae tend to develop a columel-
lar and basal callus upon cessation of growth. This has
the effect of covering over threadlike apertural bar-
riers on the columellar wall and basal lip. Much of the
variation in presence or absence of a threadlike col-
umellar barrier recorded in Table V probably is di-
rectly caused by the degree of basal callus formation. It
shows particularly clearly in the illustrations of Pal-
line biakensis (fig. 96b), P. micramyla (fig. 95d), P. no-
terapalauana (fig. 94e), and Vatusila kondoi (fig. 82b).
In complete contrast, taxa such as Kubaryiellus
kubaryi (Mollendorff, 1900) and Russatus nigrescens
(Mollendorff, 1900) (fig. 89b, e) totally lack any trace of
such a callus.
A final comment is required concerning the de-
velopment of sulci on the whorls and the very unusual
canalization of the sutural area. In the Endodontidae
(Solem, 1976b, pp. 51-52) development of a subsutural
and/or subperipheral sulcus is primarily associated
with keel formation. In the two sharply keeled Char-
opidae, Maafu thaumasius (fig. 76a-c) and Himero-
concha quadrasi (Mollendorff, 1894) (fig. 105b), quite
prominent sulci are present both above and below the
periphery. In H. fusca (Quadras & Mollendorff, 1894)
(fig. 105e), which is much less sharply keeled, only a
weak supraperipheral sulcus is present. In the one
species with angulated periphery, Sinployea angularis
(fig. 64e), there is a clearly defined subsutural sulcus,
and in both Vatusila kondoi (fig. 82b) and Sinployea
rudis (fig. 47e), which have rounded peripheries, there
are distinct supraperipheral sulci. They are the only
equivalent to the unusual sulcus formation reported in
the endodontid genus Anceyodonta (Solem, 1976b, p.
51, figs. 81d-e, 83, 86a, c, 88a, c, 89a). The functional
significance, if any, of this last development is un-
known.
A most unusual modification involves canalization
of the suture. This is more clearly seen in Sinployea
proxima (Garrett, 1872) (fig. 46a-b) where it is accom-
plished by slight detachment of at least part of the body
whorl and creation of a narrow channel at the suture.
This also occurs in Lagivala macroglyphis (Rensch,
1937) (fig. 81d-e), where the channel is formed by de-
tachment to a greater angle and thus is shallower and
wider, and to a barely detectable extent in L. micro-
glyphis (Rensch, 1937). The same phenomenon is seen
in the New Caledonian species Andre francia alveolus
(Gassies, 1881) (Solem, 1961, pp. 454-456, fig. 12) and
to a lesser extent in some Australian and New Zealand
taxa. There is no equivalent change in the Endodont-
idae.
SHELL SCULPTURE
Data on the basic method of sculpture formation,
apical sculpture type, and sculpture spacing is pre-
sented on pp. 40-41 as part of the family comparison
discussion. In general the postapical sculpture in the
Charopidae is relatively uniform in composition. It
consists of major ribs, a few microradials in between,
and fine microspiral segments that act as lateral sup-
ports to the microradials (figs. Id-e, 2b, d), much in the
same way as the buttresses to the microradials found
in Ptychodon microundulata (Suter, 1890) (Solem,
1970b, pi. 59, figs. 9-11). The latter are quite different
from the microspirals found in the Endodontidae, but
this difference can be investigated only with the SEM.
In some of the charopid species with reduced radial
sculpture, such as Himeroconcha, these microspirals
become enlarged into essentially continuous spiral
cords. Again, this is far less specialized than in some of
the situations found in the Endodontidae.
As an example of the specializations in microspiral
elements, compare Figures Id-e, 2d, and 3b, e. In
Sinployea modicella (Ferussac, 1840) (fig. Id-e) the
microspirals are on both sides of the radial riblets and
tend to be joined medially and form minor cordlets. In
FIG. 1. Shell microsculpture in Sinployea modicella (Ferus-
sac). Station 893, Faatoai Valley, Moorea, Society Islands. BPBM
150377: a-b, apical and early postnuclear sculpture, note radial
element on apex in a (295x) and secondary spiral "cording" in b
(310x); c, detail of apical sculpture, note minute radial undula-
tions and even nature of spiral cords (970 x); d, postnuclear
sculpture between two major ribs on body whorl (960 x); e, detail
of microsculpture, note uneven nature of microspiral buttresses
(2,950x).
PATTERNS OF MORPHOLOGICAL VARIATION
11
contrast, Tuimalila pilsbryi (fig. 2d) shows some of the
same kind, but in the same inter-rib area it can have
the microspirals as buttresses on only one side of the
microradial. Palline biakensis (fig. 3b, e) and Micro-
charopa mimula (fig. 4a) have very minor actual
surface microspiral elements (fig. 3e), simply a very
narrow support ridge leading up to the top of the mi-
croradial whose upper edge then expands into an elon-
gated bead (fig. 3e) upward along the riblet. These
"beads" appear, when viewed at lower magnification
(fig. 3b), to form microspiral "cords" that at optical
magnification are equivalent in aspect to the serrated
edges on the microriblets in the endodontid Austral-
donta raivavaeana Solem (1976b, p. 33, fig. 23a-d). In
both situations the spiral sculpture is barely visible at
100 x magnification, and determination of the compo-
FIG. 2. Shell microsculpture in Sinployea peasei and Tuimalila pilsbryi: a-b, Sinployea peasei. Station 14, Avana Valley, Rarotonga, Cook
Islands. FMNH 144613. a, apex and early postnuclear sculpture (320 x); b, suture between apex and first postnuclear whorl, note beaded
secondary spiral "cording" and waved effect of apical cords caused by conforming to minute radial undulations on surface ( 1,065 x); c-d,
Tuimalila pilsbryi. Station T-22, east side of Eua, Tonga. FMNH 152378. c, Apical and early postnuclear sculpture, note definite, widely spaced
large radial swellings on apex, reduced size of spiral cords (320x); d, postnuclear sculpture on body whorl, note varying height of microspiral
buttresses (l,065x).
FIG. 3. Shell microsculpture in Palline biakensis. Hospitaal-
grot, Biak, West Irian. Holotype. Rijksmuseum van Natuurlijke
Historie, Leiden: a, apical and early postnuclear sculpture (275
x); b, 1st and 2nd postnuclear whorls (280x); c, detail of apical-
1st postnuclear whorl suture (930x); d, end of apical shell
sculpture (920x); e, detail of sutural area between 1st and 2nd
postnuclear whorls (910 x).
12
FIG. 4. Shell microsculpture in Microcharopa mimula. Cen-
tral peak, Mothe, Lau Archipelago, Fiji. BPBM 78585: a, apex
and part of 1st postnuclear whorl (295x); b, suture between apex
and 1st postnuclear whorl (970x); c, center of apex (980x); d,
center of apex (2,935x); e, detail of apical ridge (9,850x).
13
14
SOLEM: ENDODONTOID LAND SNAILS
nents creating the visual effect requires l,000x SEM
viewing.
The transition between apical and postapical
sculpture is quite abrupt (fig. 3d). There is a very short
distance of stuttering apical growth, then a start of the
typical postapical sculpture immediately. No dif-
ferences in this pattern were seen in other species
examined with the SEM.
Rib counts were made on the body whorl of all
unworn adult specimens. The pattern of mean rib
counts and rib spacing is presented in Table I and also
discussed in the family comparison section. The ques-
tion of the utility of rib counts is covered by Solem
(1976b, pp. 39-44), and justification of this is not re-
peated here. Data on shell diameter and rib spacing
correlations are presented in Table III. The correlation
between shell diameter and rib spacing is less tight
than in the Endodontidae (Solem, 1976b, p. 44, table
XV), probably reflecting in part the smaller number of
species involved. Rib reduction is discussed elsewhere
(p. 9) and also is not as tightly correlated with size as in
the Endodontidae (Solem, 1976b, pp. 47-50).
In the Endodontidae and the Charopidae there are
a few situations in which the major ribs have become
grossly enlarged and widely spaced. Although none of
the charopids equals the spectacular "wings" of Zyz-
zyxdonta alata Solem (1976b, p. 466, fig. 198a-c), one
does come close to the huge ribs of Cookeconcha stell-
ulus (Gould, 1844) (Solem, 1976b, p. 218, fig. 93).
Palikirus cosmetus (fig. 91d-f) from Ponape is the one
example of spectacular rib enlargement in the
Charopidae. Even when a keel is formed and the shell
enlarged as in Maafu thaumasius (fig. 76a-c), the ribs
themselves remain relatively small. When there is
great size enlargement of the shell in such endodontid
genera as Gambiodonta (Solem, 1976b, pp. 435-442,
figs. 186-189), sometimes (G. mirabilis Solem, 1976; G.
grandis Solem, 1976) there is clear rib enlargement.
Variation in the number of apical cords between
charopid species is relatively small and generally is
TABLE III. - SHELL DIAMETER AND RIB SPACING IN THE CHAROPIDAE.
Ribs/nun.
LESS THAN 2
2.0-2.99
3.0-3.99
4.0-4.99
5.0-5.99
6.0-6.99
7.0-7.99
8.0-9.99
10.0-12.99
13.0-19.99
MORE THAN 20
Number of
Taxa
2
1
2
5
4
6
5
14
26
14
6
Mean and Range
of Shell Diameter
in mm.
6.06(4
2.40
4.00(3
4.88(3
3.29(2
3.45(2
2.69(1
3.06(1
2.61(1
2.29(1
1.91(1
59-7.52)
29-4.70)
49-5.68)
11-4.30)
17-5.23)
75-3.49)
65-4.63)
73-3.74)
40-2.95)
07-3.21)
greatly exceeded by the variation within species. A
range of four to six within a population or species is not
unusual where the mean is 11 or 12, dropping to a
range of three to four where the mean is eight or nine
cords. Such variation in part reflects accidents of post-
nuclear whorl attachments that can change the count
by one or two cords, which is partly what happens in
terms of cord fragmentation near the upper suture and
is partly actual variation in cord width and spacing.
Because the intrapopulational variation is so large, lit-
tle importance is attached to minor mean differences.
Data on 83 of the 95 taxa were available. Of the re-
maining 12, Discocharopa has only radial apical
sculpture, three had an apical sculpture too fine to
count optically, five had the apex too worn, and three
had the apical whorls missing or were worn fossils.
Table I presents the dispersal of the species means,
which shows a rather tight central cluster. Of the five
species with a mean of less than 8.5 spiral cords, three
(Roimontis tolotomensis, Himeroconcha lamlanensis,
and Kubaryiellus kubaryi) have a moderately de-
pressed to sunken spire, Jokajdon callizonus (Mollen-
dorff, 1900) (fig. 92a-c) may be involved in secondary
size reduction, and only Vatusila vaitupuensis (fig.
85a-c) shows no special features explaining the reduc-
tion in number of apical cords. At the other extreme,
Lauopa mbalavuana (fig. 77a-c) with 22 cords, Mi-
crocharopa mimula (fig. 4a) with a mean of 17.8,
Sinployea irregularis (Garrett, 1872) (fig. 65d-f) with
18.5, S. recursa (fig. 64a-c) with 15.9, S. angularis (fig.
64d-f) with 14.6, and Palline biakensis (fig. 96a-c)
with 14.0 cords show no clear indication of why the
numbers have increased. Even the very elevated spire
of Ba humbugi (fig. 74b) did not make a noticeable
difference in apical cords (mean 12.0). The number is
undoubtedly much higher in the species of Tuimalila
(fig. 2c), but these cords are nearly suboptical in size
and are partly obscured by the radial swellings.
Variation in the height of the apical cords can be
studied easily with the SEM. A few examples are
shown in Figures 1-3. The situation in Sinployea
modicella (Ferussac, 1840) is typical (fig. la-c), with
the height of each cord (mean 9.2 cords) being slightly
less than its width. There is an obscure radial rugosity
as a secondary feature. Within the same genus,
Sinployea peasei (fig. 2a-b) (mean 11.4 cords) shows a
reduction in both height and width of each cord, al-
though the underlying rugosity is just as clearly out-
lined. In Tuimalila pilsbryi (fig. 2c) the spiral cords are
about the same size as in Sinployea peasei, but the
secondary radial swellings confuse the viewer using
optical equipment. Finally, in Palline biakensis (fig.
3a-d) the spiral cords are much higher and narrower,
and there is no trace of the radial rugosities seen in the
Sinployea.
Only two Pacific Island charopids show a different
apical sculpture. Microcharopa mimula (fig. 4a-e) has
spiral apical ribbing, but this is broken up into a series
of short segments with peculiar open ends (fig. 4d-e).
PATTERNS OF MORPHOLOGICAL VARIATION
15
At optical magnifications the sculpture appears to be
simple spiral cords, but obviously it is much more com-
plex. The same sculpture is found in the North Ameri-
can Radiodiscus millecostatus (Pilsbry & Ferriss, 1906)
(Solem, 1977b, p. 152, figs. 7-8) and an unnamed
Western Australian genus (Solem, unpublished data).
These genera are grouped in the charopid subfamily
Rotadiscinae H. B. Baker (1927, p. 228). Discocharopa
aperta (Mollendorff, 1888) (fig. 5a-c) has an apical
sculpture of fine radial ribs that become very crowded
(fig. 5a) near the end of the apex. Higher on the apex
(fig. 5b-c) there are irregularly spaced periostracal
spiral folds that add a partial spiral element. The post-
apical whorls (fig. 5d-f) show no trace of any spiral
sculpture.
In summary, the Pacific Island Charopidae mostly
have a very characteristic and conservative pattern of
sculpture. The apex has a variable number of spiral
cords to which radial swellings are occasionally added.
The postapical whorls have major radials whose spac-
ing and number are somewhat correlated with shell
diameter, finer microradials in between, with very fine
microspiral units that buttress or connect the microra-
dials. Most of this sculpture is formed by the perio-
stracum (p. 137). Reduction in prominence or loss of
sculptural features is an apparently rare event.
The situation is much more complex in relation to
the New Zealand and Australian charopid taxa.
APERTURAL BARRIERS
Traditionally, those endodontoid taxa with aper-
tural barriers were placed in the form genus En-
dodonta, and those without barriers, in the form genus
Champa (Solem, 1976b, pp. 118-119). The discovery
that taxa with apertural barriers showed family-level
differences in anatomy is one of the more significant
results of this study. Solem ( 1973b) reviewed the bar-
rier frequency and structure in the two families and
concluded that although the endodontoid barriers are
of common origin, those found in the Pacific Island
Charopidae have evolved at least four times. Thus, in-
stead of considering monophyletic variation, as in the
equivalent discussion of the endodontid barriers
(Solem, 1976b, pp. 52-72), an obviously polyphyletic
situation exists.
Such barriers are a common phenomenon in many
land snail families (Solem, 1972c), but the timing of
their appearance in ontogeny, methods of formation,
state in adulthood of the individual, positions in the
aperture, length, and numbers differ radically. They
are analogous rather than homologous developments.
Retention of specialized terms for the different
structures found in separate families is fully justified. I
have chosen to use a uniform terminology for the bar-
riers in the Endodontidae and Charopidae, despite
their independent origins, for ease in comparisons and
because the barriers have the same patterns of change
and growth. In both families the barriers appear at or
very shortly after hatching, are added to anteriorly and
resorbed posteriorly during subsequent growth, and
often effectively narrow the aperture in adulthood.
Recognition of parietal, columellar, and palatal
zones in the aperture, the differences between major
barriers and small traces, and the numbering sequence
[parietals (upper to lower), columellars (upper to
lower), palatals (lower to upper), all in numerical se-
quence regardless of possible homology] have been
given by Solem (1976b, pp. 52-54, figs. 36-38).
The occurrence of barriers in the Charopidae is
neither taxonomically nor geographically random: 32
(33.7%) of the 95 taxa have parietal and 27 (28.4%)
have palatal barriers. 1 These are clustered in 11 of the
20 genera. Lauopa, Discocharopa, Ladronellum, and
Roimontis are monotypic, Jokajdon and Palikirus have
two species, Graeffedon and Palline have three, Sem-
perdon has five, and Lagivala and Vatusila have six
species. No species in these genera are without bar-
riers, except for Palikirus ponapicus (Mollendorff,
1900), which is known from a single, fungus-covered
specimen and was associated with Palikirus as a con-
venience, and great secondary reduction of the barriers
in Vatusila niueana (fig. 83e). Five of the 32 species are
basically extralimital: Lagivala macroglyphis (Rensch,
1937) and L. microglyphis (Rensch, 1937) from New
Britain; L. demani (Tapparone-Canefri, 1883) from
Ambon, Timor, and parts of West Irian; Palline biaken-
sis from West Irian; and Discocharopa aperta (Moll-
endorff, 1888), which ranges from the Philippines to
Society Islands. The three Graeffedon are limited to
Western Samoa; the Lau Archipelago has two species
(Vatusila kondoi and V. nayauana) on Nayau and
Lauopa mbalavuana on Vanua Mbalavu and two
species on Viti Levu (Lagivala vivus and L. minus-
culus). Vatusila tongensis on Eua, Tonga; Lagivala
davidi (Ladd, 1968) fossil on Funafuti and Vatusila
vaitupuensis live on Vaitupu, Ellice Islands; V.
niueana on isolated Niue Island; and V. eniwetokensis
(Ladd, 1958) fossil on Eniwetok in the Marshall Islands
complete the western distribution. A second cluster of
taxa occurs on some of the Caroline, Palau, and
Mariana Islands. On Ponape there are Palline mi-
cramyla, Jokajdon callizonus (Mollendorff, 1900), J.
tumidulus (Mollendorff, 1900), Palikirus cosmetus
(Mollendorff, 1900), and Roimontis tolotomensis. On
the islands of Palau, the three subspecies of Palline
notera, Semperdon uncatus, S. xyleborus, and S.
kororensis (Beddome, 1889), are found. In the Mariana
Islands S. heptaptychius (Quadras & Mollendorff,
1894) and Ladronellum mariannarum (Quadras &
Mollendorff, 1894) live on Guam, and Semperdon
rotanus occurs on Rota and the northern tip of Guam.
'This differs from the figure cited by Solem (1973b, p. 301) be-
cause Discocharopa aperta (Mollendorff, 1888) has some populations
with and many without a parietal barrier, so this species was omitted
from that calculation.
FIG. 5. Details of shell sculpture in Discocharopa aperta (Mollendorff). Busuanga, Philippine Islands. FMNH 18725: a, shell sculpture at
apical-postnuclear boundary (l,000x); b, apical and postapical sculpture (l,035x); c, details of apical sculpture (3,000x); d, postnuclear
sculpture (l,015x); e, postnuclear sculpture showing serrated edging (3,000x); f, detail of postnuclear sculpture showing absence of spiral
elements (9,950x).
16
PATTERNS OF MORPHOLOGICAL VARIATION
17
Except for the scattered occurrence of Dis-
cocharopa aperta (Mollendorff, 1888) in the Austral Is-
lands and on Borabora, Society Islands, the entire Cook
and Society Islands area of great charopid diversity is
free from barrier-bearing genera. Only the one taxon is
known from Tonga, although several Sinployea and
Tuimalila live in this group. Both the Lau Archipelago
and Viti Levu have more taxa without than with bar-
riers, as does Samoa. In Micronesia, Guam has the en-
demic barrierless Himeroconcha, and Ponape has two
genera, Kubaryiellus andRussatus, that lack barriers.
The Palau group is unique in having only barrier-
equipped taxa in both the Endodontidae and Charo-
pidae. On some other Caroline Islands, there are
barrier-free taxa, and on both Funafuti and Vaitupu
there are species of Sinployea. The extension of
barrier-equipped taxa to the Marshall Islands repre-
sents their only penetration beyond the barrier-free
taxa.
A summary of the evidence that the barriers in
these genera represent independent derivations was
given in Solem (1973b). The uniform pattern of micro-
armature and growth in the Endodontidae was re-
viewed both in Solem (1973b) and in more detail sub-
sequently (Solem, 1976b, pp. 54-66, figs. 39-42). In
brief, the tops of the endodontid barriers normally are
modestly expanded into a cordlike shape, with very
fine triangular points (Solem, 1976b, figs. 39e, 40,
41a-c, 42a-f) that are added to the surface. They are
not parts of basic crystal layers that simply extend
outward, but are an extra feature. Checking the re-
sorption edge of a barrier (Solem, 1976b, p. 64, fig. 40)
shows that they, at times, will be covered up by sub-
sequent growth as the height of the barrier is in-
creased.
The patterns in the Charopidae are only partly
studied. At the time that most of the material used was
on loan for study (1962-1967), the SEM was not yet
available. In some of the genera, Lagiuala, Vatusila,
and Graeffedon, specimens are few in number and the
barriers too deeply recessed for adequate SEM study
without breaking the shell. Thus the following com-
ments must be viewed as only introductory.
In Jokajdon callizonus (Mollendorff, 1900) (fig.
6a-d) from Ponape, views of the third palatal barrier
demonstrate the basic structures in one species com-
plex. In vertical view of the posterior descending face,
crest, and upper anterior descending face (fig. 6a), the
etched resorption plane is on the left, whereas the right
shows the larger platelets on the upper anterior slope
of the barrier. At higher magnification (fig. 6b), the
mixture of large and small platelets on the upper an-
terior face is seen in vertical view. In Figure 6d, the
same area is shown in lateral view to demonstrate that
the platelets are simply prolonged crystals from the
growth face of the barrier. Figure 6c, at the point
where the sharp anterior descension of the barrier
ends, shows that there is an abrupt shift from large to
small platelets. At this point, the combined major func-
tion of gripping surface for extending the body from the
shell aperture and providing setal catching projection
against arthropod entry becomes much less significant,
and hence the probable shift from large to small
platelets.
Looking at the first palatal barrier in the Western
SamoanGraeffedongraeffei (Mousson, 1869) (fig. 7a-c)
shows a second pattern of structure. The specimen was
taken alive, and the shell was partly fractured during
dissection. The barrier surface was not worn, so that
the structure of long, straight, projecting ridges across
the anterior descending face of the barrier is normal.
There is distinct variation in the length of the ridges
(fig 7b-c). The most intriguing aspect is seen on the
side of the barrier (fig. 7a). The individual pockets hold
platelets at a variety of angles, sometimes protruding,
sometimes almost vertical in orientation. I consider it
possible that they represent fragments from the ad-
sorption surface that have been transported and re-
fixed in a near anterior position. Unfortunately,
specimens of Graeffedon are quite rare, and it is im-
probable that material sufficient to test this idea will
become available for study.
The Palau Island Semperdon xyleborus (fig. 8a-e)
has long parietal barriers, with the upper parietal
posteriorly bifurcate (fig. 8a). The upper edge of the
third parietal (fig. 8b-c) shows a few broad, projecting
plates that point anteriorly. Some of them stretch
across almost the entire edge. In lateral view (fig. 8d),
the edge of the fourth parietal shows that the plates
become much smaller on the side of the barrier and
that the large plates are growing from the barrier sur-
face as it slants slightly downward. They are partly
elevated from the surface.
The basic difference between the three types can
be summarized as: ( 1 ) crystal layers simply continued
directly forward and broken into microplatelets (Jo-
kajdon); (2) anterior added ridges that may be in part
transported as large crystals (Graeffedon); and (3)
broad ridges angled up from the surface layers (Sem-
perdon). Despite similar appearance and function,
these are actually three different ways of reaching the
same end result, i.e., a series of microprojections that
provide gripping surfaces for the advancing mantle col-
lar during emergence of the body from a fully with-
drawn position. They are just as effective as the fourth
solution, the triangular points in the Endodontidae,
but are not as elegantly constructed. Coupled with the
very obvious size and position differences of the bar-
riers in the three charopid examples cited, which are
typical of their anatomical groupings, I conclude that
they are of independent origin. When compared with
the apertural barrier microstructure in other families
(Solem, 1972c), the charopid experiments show greater
individual variation and are much less "finished" in
appearance, even though the use of angled plates is a
more common pattern than the addition of points or
barbs on the surfaces.
The presence of barriers in the Charopidae is a
polyphyletic and secondary character, rather than the
norm as in the Endodontidae. Thus, the comparative
SOLEM: ENDODONTOID LAND SNAILS
FIG. 6. Microstructure of largest palatal barrier mJokajdon callizonus (Mollendorff). Station 118, Ponape, Caroline Islands. BPBM 154161:
a, vertical view onto top of largest palatal at l,030x, note weak etch marks on upper left and the clear cross-strata pattern on entire posterior
(left margin); b, area of sharp anterior slope in vertical view at 3,200x; c, area at bottom of sharp anterior descension showing transition from
large crystal plates to small crystals at 3,125x; d, lateral view at 3,050x of large crystal area showing how these plates are continuations of the
vertical growth layers.
treatment will be different than in Solem (1976b).
Correlations among barrier condition and other shell
features are not as simple and direct as in the En-
dodontidae. For example, the relative degree of aper-
ture narrowing in the Endodontidae (Solem, 1976b, p.
71, table XLVIII) and Charopidae (table IV) is quite
different. The percentages of the barrier-possessing
taxa that are intermediate in closure differ greatly. In
the Endodontidae, 57.5% are intermediate, but only
25.0% in the Charopidae. The latter have the barriers
TABLE IV. - DEGREE OF APERTURAL NARROWING BY BARRIERS
Strong
Moderate
Weak
Not
Total Taxa
Endodontidae
19.9%
40.9%
16.6%
22.6%
181
Charopidae
43.7%
9.4%
15.6%
31.3%
32
PATTERNS OF MORPHOLOGICAL VARIATION
19
FIG. 7. Surface of largest palatal barrier in Graeffedon graeffei
(Mousson). Station 39, Mt. Solaua, Upolu, Western Samoa. FMNH
153420: a, angled view of front edge and inner side showing different
lateral and anterior surface texture (305 x); b, anterior edge at 1,100
x showing projecting platelets on surface of descending edge; c, ver-
tical view onto descending edge surface at 3,150x .
either strongly constricting the aperture (figs. 98b, e,
99b, d, e), or they are essentially ridgelike remnants
(figs. 85b, lOlb).
In both families, more species have parietal bar-
riers than either columellar and palatal, and no species
has either of the latter without also having the former.
Parietal barriers
The number of parietal barriers ranges from the
single barrier that may be present or absent in Dis-
cocharopa, to the four large barriers plus three traces
found in Semperdon xyleborus (fig. 98e). The only vari-
ation in major barrier numbers occurs in Semperdon
heptaptychius (Quadras & Mollendorff, 1894), which
normally has three parietals and one accessory trace.
Occasionally (5.1%), the trace and third parietal are
missing, rarely (0.3%), a fourth parietal is developed,
and rarely (3.3%), the trace is missing. In Semperdon
xyleborus, 84.6% have three traces, 15.4% have two
traces. Occasionally, a parietal trace will be present in
Palline notera notera and P. n. gianda, and there is one
superior trace present in S. rotanus (fig. 99d). Other-
wise, the number of parietal barriers is constant.
Barrier length partly correlates with height. The
method of determining length is explained by Solem
(1976b, p. 70, fig. 43). Where the parietal is a thread-
like remnant, it is Vie to Vs whorl long (figs. 83e, 85b,
95d, lOlb), except in Palline biakensis (fig. 96b) and
Roimontis tolotomensis (fig. 91b) where the length
reaches 3 /ie of a whorl. In Vatusila nayauana (fig. 82e),
Semperdon uncatus (fig. 97b), and Palikirus cosmetus
(fig. 91e), the parietal is a raised ridge, but still short.
The only taxa with high, short barriers are the races of
Palline notera (figs. 94b, e, 95a) and Semperdon
xyleborus (figs. 8a, 98e). The remaining barriers are V*
to 3 /ie of a whorl long, except for several species of
Lagivala where they extend to or beyond the line of
vision from the aperture (L. microglyphis, fig. 81b; L.
macroglyphis, fig. 81e; L. minusculus, fig. 80e). These
species are very small in size, with mean diameters of
1.55-1. 73 mm.
Structure of the barriers is highly varied. The
threadlike and raised ridge taxa present no unusual
features and have been mentioned above. Where there
is a single high barrier, such as Lauopa mbalauuana
(fig. 77b), Discocharopa aperta (Mollendorff, 1888) (fig.
35a-b), and Lagivala minusculus (fig. 80e-f), it is a
high thin blade, usually with abrupt anterior descen-
sion. When there are two or three high lamellae, the
typical shape found in most of the Endodontidae is
usual: an abrupt posterior resorption face, elevated
middle with slightly to strongly expanded upper edge,
and then gradual anterior descension. The second and
fourth parietals in Semperdon xyleborus (fig. 8a) are
typical. Several taxa have a modification of the first
parietal that also was found in some Endodontidae. It
can become deflected downward on the posterior por-
tion in Jokajdon callizonus (Mollendorff, 1900) (fig.
92b), Semperdon heptaptychius (Quadras & Mollen-
FIG. 8. Structure of apertural barriers in Semperdon
xyleborus. Station 184, Peleliu. Palau Islands: a, parietal barriers
at 120x, note posterior bifidity of upper (1st) parietal: b, upper
edge of 3rd parietal at 950x; c, upper edge of 3rd parietal at
2,850x; d, upper edge of 4th parietal at 2,910x; e, lower palatal
barriers at 326x . Anterior side at left margin in each photograph.
20
PATTERNS OF MORPHOLOGICAL VARIATION
21
dorff, 1894) (fig. 99b), and S. rota nut* (fig. 99d); it is
bifid in Vatusila tongensis (fig. 83b) or posteriorly
bifurcated in V. kondoi (fig. 82b), Jokajdon tumidulus
(Mollendorff, 1900) (fig. 92e), Semperdon kororensis
(Beddome, 1889) (fig. 98b), and S. xy/ebon/s (fig. 8a,
upper barrier). Similar bifurcation was found also in
three lineages of the Endodontidae several Cooke-
concha sand Endodonta (Solem, 1976b, p. 379, fig. 167a,
g), some Min idon ta (Solem, 1976b, p. 138, fig. 65b,d-e,
g; p. 149, fig. 71c), and some Anceyodonta (see Solem,
1976b, p. 57). The reason for these repetitive bifurca-
tions, usually of the upper parietal barrier, is un-
known.
The basic similarity in shape between the larger
parietals of the Charopidae and Endodontidae proba-
bly indicates that this is an efficient shape, not only in
terms of continual anterior addition and posterior re-
sorption, but also in functioning during body extension
of the animal.
None of the Charopidae show the splitting into
threadlike traces of the barriers that was common in
several lines of the Endodontidae (Solem, 1976b, p. 57;
p. 332, fig. 144a-c).
The Charopidae show instead a quite conservative
shape and number pattern of the parietal barriers.
Columellar barriers
Of the 32 taxa with parietal barriers, 13 lack any
columellar barrier. This part of the shell is missing in
Vatusila eniwetokensis (Ladd, 1958), so that its situa-
tion is unknown. Semperdon xyleborus and S. kororen-
sis (Beddome, 1889) (fig. 98b-c, e) have two prominent
columellar barriers, and the former species normally
has two accessory traces above the upper columellar.
Six taxa, Lagivala macroglyphis (Rensch, 1937) (fig.
81e-f), L. microglyphis (Rensch, 1937) (fig. 81b),
Jokajdon callizonus (Mollendorff, 1900) (fig. 92b), J.
tumidulus (Mollendorff, 1900) (fig. 92e-f), Graeffedon
savaiiensis (fig. 88b), and Pa/line notera notera (fig.
94b) have only one columellar. This varies from a
threadlike ridge in the last two, to high and sometimes
sharply ascending barriers in the other four species.
The remaining 10 taxa (table V) show intraspecific
variation in the number or absence of columellar bar-
riers. When the variation is present or absent, in all
but Lagivala demani (Tapparone-Canefri, 1883), the
barrier is minute and threadlike. Its absence would
mostly result from being covered by the basal callus. In
Lagivala demani (Tapparone-Canefri, 1883) the varia-
tion is geographic, with a large columellar present in
specimens from Timor, medium-sized in Ambon and
Aru Islands shells, and very small to absent in Biak
and West Irian examples. In Semperdon rotanus and S.
heptaptychius (Quadras & Mollendorff, 1894) (fig. 99e)
the second columellar, when present, is a small thread-
like trace above the larger columellar that is a high
ridge parallel to the plane of coiling. A large form of
the latter species (fig. 99b) occurs in which the columel-
lar barrier is absent.
TABLK V. - PERCKNTACK DISTRIBUTION OF COLUMKI.I.AR BARRIER NUMBERS.
Vatusila kondoi
V. nayauana
Lagivala demani
Graeffedon graeffei
Palline notera gianda
P. notera palauana
Number of Columellar Barriers
1
5.6
94.4
40.0
60.0
*
*
30.0
70.0
84.0
16.0
95.0
5.0
83.0
17.0
46.5
0.3
42.1
3.5
96.5
53.5
56.4
Semperdon uncatus
S . rotanus
^. heptaptychius
Ladronellum roar iannarum
* Geographically variable.
In most taxa the columellar barrier lies parallel to
the plane of coiling. Semperdon xylcborus and S.
kororensis (Beddome, 1889) have one barrier slanting
downward (fig. 98b, e); Jokajdon tumidulus (Mollen-
dorff, 1900) (fig. 92e), Lagivala microglyphis ( Rensch,
1937), and L. macroglyphis (Rensch, 1937) (fig. 81b,
e-f) have it twisted up toward or almost onto the
parietal wall; and in others, such as Jokajdon cal-
lizonus (Mollendorff, 1900) (fig. 92b), it is a deeply re-
cessed, triangular knob.
Thus, the 18 species of Charopidae known to have
a columellar barrier show almost the same total range
of variation reported for the Endodontidae (Solem,
1976b, pp. 57-59).
Palatal barriers
Five of the 32 taxa with parietal barriers lack
palatals Lauopa mba/auuana (fig. 77b),Discocharopa
aperta (Mollendorff, 1888) (fig. 36b), Palline biakensis
(fig. 96b), Roimontis tolotomensis (fig. 91b), and
Palikirus cosmetus (fig. 91e). Except for the Roimontis.
which has two threadlike parietals, all of them have
only a single, short lamellar parietal. Because of the
missing palatal wall in the holotype and only known
example, the palatal barrier situation in Vatusila
eniwetokensis (Ladd, 1958) is unknown. In Ladronel-
lum mariannarum (Quadras & Mollendorff, 1894) (fig.
lOlb), many Vatusila nayauana (fig. 82e), and V.
niueana (fig. 83e) there is a modest ridge to barely
detectable trace on the middle of the palatal wall.
Semperdon uncatus (fig. 97b) normally has one low
barrier on the basal lip, but in 17% of the individuals
this is missing, and Palline micramyla (fig. 95d) has
two low lamellar barriers on the basal lip. Lagivala
minusculus (fig. 80e) has a rather unique ridge with a
lateral buttress on the basal lip, and Vatusila tongensis
(fig. 83b) has a very large basal lip barrier that is
bifurcated or split into two parts plus a very unusual
22
SOLEM: ENDODONTOID LAND SNAILS
upper raised callus that is 0.25-0.36 mm. wide. Vat-
usila vaitupuensis (fig. 85b) has a midpalatal thread-
like trace and a broader ridge on the basal lip very near
the columellar margin.
The remaining species have a more typical array
of normally three to seven palatal barriers, sometimes
with additional accessory traces present. The position-
ing of these barriers varies from at the lip edge in Sem-
perdon (figs. 98b-c, e, 99b, d, e),Palline notera and its
subspecies (figs. 94b, e, 95a), Graeffedon (figs. 88b, e,
86b), and Lagivala microglyphis (Rensch, 1937) (fig.
81b), to partly recessed in Jokajdon callizonus (Moll-
endorff, 1900) (fig. 92b), progressively recessed in J.
tumidulus (Mollendorff, 1900) (fig. 92e), moderately
recessed in Vatusila kondoi (fig. 82b) and Lagivala
macroglyphis (Rensch, 1937) (fig. 81e), and deeply re-
cessed in both L. vivus (fig. 80b) and L. davidi (Ladd,
1968) (Ladd, 1958, pi. 30, figs. 13-15).
The shape of the barriers is rather variable among
these species, although constant within a particular
species. A simple crescent-shape with modestly to
moderately expanded upper edge is found in Lagivala
vivus (fig. 80b), L. macroglyphis (Rensch, 1937) (fig.
81e), andL. demani (Tapparone-Canefri, 1883); a more
expanded upper edge, in L. microglyphis (Rensch,
1937) (fig. 81b) and Graeffedon pricei (fig. 88e); typical
structure, in Vatusila kondoi (fig. 82b); very high, in
Graeffedon graeffei (Mousson, 1869) (fig. 86b) and G.
savaiiensis (fig. 88b); typical, in the races of Palline
notera (figs. 94b, e, 95a); and bulbous-edged, again in
Semperdon rotanus and S. heptaptychius (Quadras &
Mollendorff, 1894) (fig. 99b, d-e). In Semperdon xyle-
borus (figs. 8e, 98e), the crescent is anteriorly elon-
gated, a process continued in the two species of Jokaj-
don (fig. 92b, e,) and reaching the typical parietal bar-
rier shape in Semperdon kororensis (Beddome, 1889)
(fig. 98b).
The predominance of the crescent-shaped palatal
barriers outlined above is very different from the situa-
tion in the Endodontidae. In that family (Solem, 1976b,
p. 60, table XXXII) only five species had crescentic
palatals, whereas 130 had bladelike barriers. In the
Charopidae, only the two Jokajdon, Semperdon koro-
rensis (Beddome, 1889), and possibly S. xyleborus could
be considered to have a bladelike palatal barrier form.
Intraspecific variation in the number of palatal
barriers is summarized in Table VI. The great varia-
tion found in the two Semperdon exceeds that found in
any of the Endodontidae (Solem, 1976b, p. 67, table
XL). None of the ratios in Table VI convincingly
suggests genetic dominance. Assigning the most fre-
quent percentage to represent that species, the pattern
of palatal barrier numbers is summarized in Table VII.
It is evident that there is considerable variation within
genera, particularly Lagivala, Graeffedon, Palline, and
Semperdon. Even within genera, there is no clear cor-
relation between number of barriers and shell size.
Only Palline notera shows a partial correlation, in that
the two larger subspecies have fewer and smaller-sized
TABLE VI. - PERCENTAGE DISTRIBUTION OF PALATAL BARRIER NUMBERS
Taxon Number of Palatal Barriers
012345
Vatusila kondoi 11.1 88.9
V. nayauana 40.0 60.0
Lagivala demani *
20.0 80.0
10.0 75.0 10.0 5.0
16.0 68.0 16.0
1.2 3.0 19.8 55.0
1.2 80.0 13.1 3.9 1.5
92.0
21.0
0.3
8.0
Palline notera
palauana
V_. ri. gianda
Semperdon uncatus 17.0 83.0
S. rotanus
S. heptaptychius
. xyleborus
* Geographically variable.
palatals than the nominate race, which averages
0.24-0.41 mm. less in diameter than the others. There
is also no clear correlation between number of palatals
and degree of apertural narrowing.
The presence of palatal traces is sporadic. Several
taxa will have one or two traces between lower pairs of
palatal barriers: Lagivala microglyphis (Rensch, 1937),
Graeffedon graeffei (Mousson, 1869), G. pricei, Palline
notera gianda, Semperdon rotanus, and S. hepta-
ptychius (Quadras & Mollendorff, 1894). Other taxa will
have three traces above the upper palatal: Jokajdon
callizonus (Mollendorff, 1900), Semperdon kororensis
(Beddome, 1889), and most examples of S. xyleborus. A
few of the latter will have four (7.5%) or five (7.5%)
traces above. Jokajdon tumidulus (Mollendorff, 1900)
normally has three traces above the upper palatal and
three (rarely two) between lower pairs. The same pat-
TABLE VII. - PALATAL BARRIER NUMBERS IN CONSTRICTED APERTURE CHAROPIDAE.
3 Palatals
Vatusila kondoi
Lagivala demani
Jokajdon callizonus
J_. tumidulus
4 Palatals
Lagivala davidi
Graeffedon pricei
Palline notera gianda
P_. _n. palauana
Semperdon heptaptychius
5 Palatals
Lagivala microglyphis
U macroglyphis
Graeffedon graeffei
Palline notera notera
6 Palatals
Lagivala vivus
7 Palatals
Graeffedon savaiiensis
Semperdon rotanus
S. xyleborus
S. kororensis
PATTERNS OF MORPHOLOGICAL VARIATION
23
tern of traces above the upper palatal is seen in the
Endodontidae (Solem, 1976b, p. 59) and was inter-
preted as a space-preserving function, because the
great size of the first parietal and the need to keep this
space open for withdrawal and extension of the buccal
mass and foot combine to make reduction in size of
palatal projections in this area advantageous.
As in the Endodontidae, the normal pattern of size
change is for the palatal barriers to decrease in height
as they ascend the palatal wall. An obvious exception
will be reduced size of the first palatal in situations
where it is right at the columellar margin (fig. 99d) or
where there is large size to the columellar and/or lower
parietal barriers (figs. 86b, 92b, e). So few taxa show
either recession of the palatals or major upper edge
expansion of the barriers that discussion of shell corre-
lations separately is not attempted. Most of the com-
ments made about general features of the palatal bar-
riers in the Endodontidae (Solem, 1976b, pp. 59-63)
seem to apply to the few Charopidae also.
Despite only 27 charopids having palatal barriers,
they show a wide range of variations that mostly paral-
lel the situations found in the Endodontidae. The
biggest difference lies in the basic shape of the bar-
riers: bladelike in 86.1%, ridged in 10.6%, and
crescent-shaped in 3.3% of those Endodontidae with
palatal barriers compared with bladelike in 15.4%
(four), a lamellar or unusual ridge in 23.1% (six), and
crescent-shaped in 61.5% (16) of the 26 charopid taxa
for which data on the palatal barrier structure were
available. Judging from the size of the parietal barriers
(fig. 85e), Vatusila eniwetokensis (Ladd, 1958) probably
had large palatal barriers, but the entire outer wall of
the only known specimen is missing, and thus the state
of the barriers is unknown.
Patterns of barrier variation
Despite the lack of size and shape correlations
with variation in the barriers found in the Charopidae,
a few comments can be made concerning general pat-
terns within a particular group. In Lagivala (figs.
80-81) the variation is geographic, with the Bismarck
Archipelago species having the strongest apertural
barriers, progressive reduction occurring in both L.
demani (Tapparone-Canefri, 1883) to the west and in
the Fijian and Funafuti taxa to the east (fig. 80b, e).
Vatusila shows no size correlation, with the largest (V.
tongensis, fig. 83b) and one of the smaller (V.
eniwetokensis, fig. 85e) having quite large barriers,
and both large (V. vaitupuensis, V. niueana, figs. 83e,
85b) and small (V. nayauana, fig. 82e) having reduced
barriers. There does appear to be some geographic con-
cordance, with Tonga and Lau species having larger
barriers than taxa from the more isolated Vaitupu and
Niue localities. The fossil record of a large-barriered
form on Eniwetok in the Upper Miocene gives an his-
torical perspective to the group that otherwise is lack-
ing in the Charopidae.
The reasons for the reduction in barrier size in the
races of Palline notera on Koror (palauana, fig. 94e)
and Babelthuap (gianda, fig. 95a) compared with the
nominate race on Peleliu (fig. 94b) is unknown, al-
though slight size differences are involved. Barrier size
variation in Semperdon shows no correlations that I
could detect. The unconstricted aperture of the mean
diameter 2.83 mm. S. uncatus (fig. 97b) contrasts with
the highly constricted aperture in the mean diameter
2.88 mm. S. xyleborus (fig. 98e) and mean diameter
4.59 mm. S. kororensis (Beddome, 1889) (fig. 98b). All
of these species have been dissected and all occur on
Koror. The differences in barrier prominence cannot be
explained at present.
Summary of barrier variations
Despite the different microstructure on the barrier
edges and the evident polyphyletic origin of the aper-
tural barriers in the Charopidae, in general they paral-
lel those of the Endodontidae. The retention of parietal
barriers extending to the lip edge, general shape, and
multiple bifurcations of the parietals all suggest paral-
lel function. Given the similar habitat and overlapping
shell size, plus the same basic growth structure, this is
not surprising. The difference in shape frequency of the
palatal barriers, a greater tendency for loss or reduc-
tion in the barriers, and the general shorter length of
the charopid barriers are real average differences but
cannot be used as criteria for family-level separation
because there is so much overlap between families.
SUMMARY OF SHELL VARIATIONS
With few exceptions the range of variation in the
shells of the Charopidae is less than that found in the
Endodontidae. Major differences between the two fam-
ilies are reviewed on pp. 40-44. Here it is sufficient
to point out that the formation of sculpture by the
periostracum rather than the calcareous layers, lack of
umbilical and growth modifications to form a brood
chamber, and secondary derivation of apertural bar-
riers in the Charopidae have had profound effects on
the range of shell variation.
GROSS ANATOMY
Including the Punctum sp. from Tahiti, Sinployea
sp. from Borabora, and Sinployea sp. from Saipan,
there are 98 species-level taxa considered in this
monograph. Of these, 51 (52.0%>) belong to one genus,
Sinployea. It was possible to see at least fragmentary
soft parts for 43 (43.9%) of the total taxa and 21 (41.1%)
of the Sinployea. The taxonomic distribution of the dis-
sected material is given in Table VIII. Of the 21 gen-
era, six genera, five of them monotypic, could not be
dissected. Only Lagivala, with six species, represents
an undissected speciose group. The taxonomic coverage
of dissected material is broader than that given the
Endodontidae (Solem, 1976b, p. 73, table L), where
only 58 (32.4%) of the 179 taxa seen were dissected.
This reflects both a lesser degree of extinction for the
24
SOLEM: ENDODONTOID LAND SNAILS
TABLE VIII. - PHYLETIC REPRESENTATION OF DISSECTED TAXA.
Total
Taxa
Number
Dissected
Family Punctidae
Punctum
Family Charopidae
1
1
51
1
1
1
2
6
6
3
1
1
1
1
2
2
5
5
1
A
98
21
1
1
1
1
1
1
1
2
2
5
1
3
Charopidae and a difference in areas of species abun-
dance.
In part, the apparent greater amount of anatomi-
cal data for the Charopidae is illusory because many
taxa were represented only by pulled fragments. This
reflects the method of processing and specimen storage
used at the BPBM (see Solem, 1976b, p. 19). Most fre-
quently, a break occurs so that the digestive gland and
ovotestis do not pull out of the shell, but sometimes
only the head and terminal genitalia will be extracted.
Thus, in only 21 of the 43 taxa do I have data on the
ovotestis structure. For 14 additional taxa I could re-
cord data on the postapical genitalia and the entire
pallial region. For four taxa most of the terminal
genitalia was extracted, but only part or none of the
pallial region, and for four taxa it was possible to re-
cord data on penial structure only. Only partial review
of the anatomical patterns is possible.
The sequence of discussion follows that used for
the Endodontidae (Solem, 1976b, pp. 72-99). Expla-
nations and data presented there are not repeated, al-
though fully cross-referenced. In certain cases, refer-
ence is made to extralimital taxa that have bearing on
subfamily assignments. Where appropriate, their
anatomy is illustrated here to provide standards
against which to measure the variations found in the
Pacific Island taxa. Illustration of the anatomy for new
Australian and Lord Howe Island taxa will be pre-
sented elsewhere, although data from them are used to
establish subfamily limits.
GENITAL SYSTEM
The organs are discussed in approximate order
from apex of the coiled visceral hump to the atrium.
The terminology is modified from that originated by H.
B. Baker (1938b, pp. 6-10, 92) and follows exactly the
usage in Solem (1976b) with a few additions for new
structures.
OVOTESTIS (G) The hermaphroditic gland or
ovotestis in the Charopidae normally consists of one or
two clumps of palmately clavate alveoli that usually lie
buried in the digestve gland above the stomach-
intestine junction. The only Pacific Island species that
departs from this pattern is Discocharopa aperta (M611-
endorff, 1888) (fig. 34a), where the bilobed ovotestis
(G) lies alongside the stomach rather than apically in
the digestive gland. In Sinployea (figs. 39b, g, 43c, 53g,
57b, 60b) there may be one or two lobes, basically de-
pending on exactly where the last branching of the
hermaphroditic duct occurs. Generally the branching
of the ducts is simple, but in Graeffedon graeffei (Mous-
son, 1869) (fig. 87d) there are lateral short branches off
the individual alveoli after the initial bifurcation. In
Ba humbugi (fig. 75c) the ovotestis is shortened consid-
erably. Jokajdon, Kubaryiellus, andLadronellum have
one clump, Palline and Trukcharopa, two clumps
typical differences are seen in Figure 90f, i. Species of
Semperdon (fig. lOOc, g) have one or two clumps. In
every case these clumps lie essentially parallel to the
plane of coiling and extend apically with only an initial
angling outward from the parietal wall where the start
of the hermaphroditic duct lies.
This is a very different pattern from the perpen-
dicular to slanted orientation and division into a larger
number of lobes seen in the Endodontidae (Solem,
1976b, p. 74, fig. 44). Similar orientation and multipli-
cation of ovotestis lobes is found in some Australian
and New Zealand Charopidae, but among the Pacific
Island Charopidae and Endodontidae genera the ovo-
testis differences are diagnostic.
HERMAPHRODITIC DUCT (GD) As in nearly
all pulmonates, this duct serves to transport sex prod-
ucts from the mass of the ovotestis to the fertilization
area below the expanded stomach. Its length is directly
correlated with the length of the stomach, which, in
turn, is dependent upon the whorl count and cross-
sectional whorl area in this part of the visceral hump.
In Ba humbugi, which averages only 3% whorls and
has a comparatively wide aperture (fig. 74b), the her-
maphroditic duct (fig. 75c) is very short compared with
most other taxa. Whether the ovotestis starts after
union of the ovotestis collecting tubules with a sudden
or gradual expansion appears to be possibly seasonal.
In no species did I have samples from the same popula-
tion taken in different months of the year available in
order to check whether swelling is a sign of reproduc-
tive activity. None of the Charopidae show the "kink-
ing" of the hermaphroditic duct seen in the Endodont-
PATTERNS OF MORPHOLOGICAL VARIATION
25
idae (Solem, 1976b, p. 75, fig. 45). The charopids do
show a characteristic purple-red, iridescent sheen to
the hermaphroditic duct that is not seen in the en-
dodontids.
TALON (GT) and CARREFOUR (X) The talon
(GT) varies from a simple swelling just above the en-
trance of the hermaphroditic duct (GD) (fig. 39e) to a
circular head on a stalk (fig. 103e). The carrefour (X) is
a slight swelling in the tract after reflexion of the duct.
The same pattern holds in all the species examined
from the Pacific Islands. As usual in both the
Charopidae and Endodontidae, there is a partial reflex-
ion of the hermaphroditic duct before it enters the car-
refour. This permits the compaction of apical organs
when the animal has retracted into the shell. The gen-
eral pattern of a circular head on a short stalk in the
Pacific Island charopids contrasts with the more slen-
der head pattern in the endodontids (Solem, 1976b, p.
85, fig. 49a; p. 373, fig. 164b; p. 471, fig. 199c), but the
differences are small in many cases, and the variation
in extralimital charopids is extensive. In both families
the collecting ducts of the albumen gland empty into
the carrefour below the level of the hermaphroditic
duct entrance (fig. 39e; Solem, 1976b, p. 471, fig. 199c).
No histological studies have been made of structures in
this area.
ALBUMEN GLAND (GG) The distance between
the apex of the pallial cavity and the base of the
stomach is less in the Charopidae than in the Endodont-
idae, since the median whorl counts are, respectively,
4Va and 5V2 + . In the Endodontidae, there is more of a
tendency for the albumen gland to be elongated and to
occupy only part of this area. In the Charopidae, the
albumen gland is almost shapeless, roughly globular,
with the surface deeply indented by intestinal loops,
head of the spermatheca (S), esophagus, arteries, and
stomach base. Rarely does the albumen gland dissect
out intact, and in order to check the shape of the talon
and carrefour plus the point of entrance for the her-
maphroditic duct, some of the albumen gland tissue
has to be teased away before illustration. Thus, only
rarely can the actual shape be shown. The size of the
alveoli appears larger in the Charopidae than in the
Endodontidae, but no quantification of this was at-
tempted.
PROSTATE (DG) and UTERUS (UT) In the En-
dodontidae (Solem, 1976b, pp. 77-78; p. 373, fig. 164d;
p. 471, fig. 199c) the prostate and uterus are completely
separate tubes that are only lightly bound together by
connective tissue. In the Charopidae they are partly
(some Rotadiscinae) to completely (most Charopinae)
fused, sharing a common lumen, with the prostate
channel a lateral outpocket (fig. 102e) into which enter
the prostatic alveoli. These are much fewer in number
than in the Endodontidae, longer, and not arranged in
distinct rows. Generally the prostate is as long as the
expanded uterine upper section and extends slightly
down along the even more expanded uterine lower
chamber. The latter tapers into the free oviduct (UV)
without clear demarcation, so the exact relationship
between prostate and uterine length remains to be de-
termined by histological studies.
The uterus has a clear separation into an upper
(UT 1 ) thin-walled narrower chamber and a lower (UT 2 )
thick-walled, more expanded chamber (fig. 43c-d).
Presumably, actual egg encapsulation takes place in
the lower chamber. This is simpler than the situation
in the Endodontidae (Solem, 1976b, p. 449, fig. 191b)
where four sections could be distinguished in the
uterus. Comparative studies on the function and his-
tology of this area in both families would be quite
worthwhile. Solem (1972b, pp. 108-112) reviewed the
pattern of fused versus separated pallial gonoducts in
the Pulmonata and concluded that the separated condi-
tion found in the Endodontidae was more primitive
than the fused condition found in the Charopidae and
many European-North American families.
TERMINAL MALE GENITALIA The organs
that may be present in this complex are the vas defer-
ens (VD), epiphallus (E), penial retractor muscle (PR),
penis (P), penis sheath (PS), atrium (Y), and internal
structures of the epiphallus and penis proper. The lat-
ter regions are used in species recognition by the
snails, so that highly significant variations can be
found within genera, particularly under conditions of
sympatry. The discussion of variations in the Endodont-
idae (Solem, 1976b, pp. 78-83) should be read as
background information on the general patterns before
attempting to deal with the complexities in the
Charopidae, where few of these organs are constant in
shape.
The vas deferens (VD) starts from the end of the
prostate alveoli attachment and tapers for a variable
distance before becoming a narrow tube that reflexes
at the penioviducal angle to ascend the penis. The area
of tapering can be very short as in Russatus (fig. 90b),
long in Ladronellum (fig. 102a), extend nearly to the
penioviducal angle as in Tuimalila (fig. 79b), or highly
variable as in Sinployea (fig. 67a, e-f, h). The ascend-
ing arm of the vas deferens is without unusual features
until it reaches the next structure, which differs from
subfamily to subfamily of the Charopidae.
In the Charopinae the vas deferens (VD) passes
into an epiphallic enlargement that may be apicad or
anterior of the penial retractor muscle insertion. In
Charopa coma (Gray, 1843) (figs. 9b, d, 10) there is an
enlargement of the vas deferens about one-third of the
way from the penial retractor muscle insertion. Inter-
nally (fig. 10) it can be seen that the actual differentia-
tion occurs some distance after the swelling has begun.
Internal pilasters of the vas deferens (VD) enlarge,
terminating in finger-like projections that surround a
central cavity. Below this point, which is the start of
the epiphallus, two high pilasters and a lower ridge
line the inside of the epiphallic chamber down to the
point of entry into the penis chamber (fig. 9d). The
IE
PC
FIG. 9. Anatomy of Charopa coma (Gray). Waiwera-Puhoi Road, north of Auckland, North Island, New Zealand. A. Solem! 11-10-1962.
FMNH 135420: a, left side of foot showing size relation to shell; b, genitalia; c, details of pallial genitalia; d, internal structures of penis and
lower female tract; e, pallial region; f, details of pneumostomal area; g, bottom and top views of anterior digestive system. Scale lines as marked.
(PS).
26
PATTERNS OF MORPHOLOGICAL VARIATION
27
FIG. 10. Interior of lower vas deferens and epiphallus in Charopa
coma (Gray). Waiwera-Puhoi Road, north of Auckland, North Island,
New Zealand. A. Solem! 11-10-1962. FMNH 135420. Greatly en-
larged. (NB).
penis retractor muscle (PR) inserts as a U-shaped fan
around the lower part of the epiphallus (E). Various
modifications occur extralimitally in such taxa as
Phenacohelix pilula (Reeve, 1852) (fig. 11). Here the
penial retractor muscle inserts on the head of the
epiphallus, and the vas deferens enters almost later-
ally. The shift in muscle attachment is relatively
insignificant, but does make external recognition of
the epiphallus presence difficult if not actually impos-
sible.
The Pacific Island Charopinae mostly show, in-
sofar as it could be checked, a specialized vas defer-
ens-epiphallus junction (figs, 57e, 79c). Exceptions are
found in Graeffedon and the provisionally assigned
Discocharopa (see below). A circular sphincter sur-
rounds the epiphallic pore (DE), and a Y-shaped plug
extends through the pore to attach onto the wall of the
epiphallus as a circular ridge. In general, the walls of
the epiphallus are without sculpture or have the typi-
cal ridges (fig. 87c) seen in Charopa coma (Gray, 1843)
(fig. 10). Because of size and preservation problems,
only a few of the species could be checked for this
structure, and illustrations are given only for
Sinployea complementaria (Mousson, 1865) (fig. 57e)
and Tuimalila pilsbryi (fig. 79c). It was observed in
several other Sinployea. Normally the head of the
epiphallus expands abruptly, indeed it can be bulbous
(fig. 67a), so that external recognition is easy.
In the Semperdoninae (figs. lOOb, e-g, 102a, 103a,
c-d), the vas deferens remains a thin tube during its
ascent and enters laterally into the epiphallus, with
the penial retractor muscle inserting directly onto the
head of the epiphallus. The exact point of entry of the
vas deferens into the epiphallus differs. In Himero-
concha and Ladronellum the entry occurs slightly
below the rounded head (fig. 102d), whereas in Sem-
perdon the entry is right next to one edge of the penial
retractor muscle insertion (fig. lOOi). The epiphallus is
a double-walled tube, sometimes coiled within the
muscle sheath. The outer wall is a thin muscle sheath,
and the inner a thick glandular tube usually rolled
inward on one side.
In the Trukcharopinae (figs. 90b-d, f-g, i-j, 93b-c,
f, h) the vas deferens enters directly into the penis
through a simple pore. In all examined genera except
Jokajdon (fig. 93b-c) the vas deferens passes through
the muscle fan before entering the penis. In Jokajdon
the muscle insertion is directly on the head of the
penis, and the vas deferens enters lateral to the inser-
tion. This is interpreted as a secondary modification
probably resulting from reduction in size. The Truk-
charopinae thus have no trace of an epiphallus.
In the Rotadiscinae (H. B. Baker, 1927, pi. 16, fig.
14; pi. 17, figs. 22, 27) the epiphallus is less clearly
differentiated externally, and it is either before
(Radioconus) or after (Rotadiscus, Radiodiscus) inser-
tion of the penial retractor muscle. The enlarged pilas-
ters of the epiphallus are the main differentiating fea-
tures.
In all of the Pacific Island Charopidae examined to
date the penial retractor muscle originates from the
diaphragm near the apex of the pallial cavity and in-
serts onto the penis or epiphallus. There is no shift to
the columellar muscle equivalent to that seen in the
Endodontidae (Solem, 1976b, pp. 81-83), which per-
mitted elongation of the penis in such taxa as En-
dodonta and Australdonta. In addition, generally the
penial retractor muscle in the Charopidae is very
short, at times scarcely a tuft connecting the dia-
phragm and penis or epiphallus head. This is another
correlative of the reduced whorl count in the
Charopidae as compared with the Endodontidae.
With a few exceptions, the external appearance of
the penis in the Pacific Island Charopidae is for a bul-
bous apical section that then tapers either abruptly or
quickly to a slender stalk of variable length that joins
the vagina to form the short atrium. The degree of the
apical bulge depends directly upon the size and com-
plexity of the internal pilasters and vergic structures.
Figure 67 gives a fair sample of shape variations, and
KD
GD
FIG. 11. Anatomy of Phenacohelix pilula (Reeve). Church Road, Kaitaia, Northland, North Island, New Zealand. L. Price! X-1962. FMNH
135421: a, pallial region; b, extended foot showing caudal horn (CH) and pedal grooves (FS); c, genitalia; d, junction of hermaphroditic duct (GD)
and talon (GT); e, internal structure of penis. Scale lines as marked. (PS).
28
PATTERNS OF MORPHOLOGICAL VARIATION
29
this is within a single genus. The major exception is
the Semperdoninae (figs. 100, 102, 103) where a penis
sheath, narrowed collar between epiphallus and penis,
plus a quite different pattern of internal structure have
combined to produce a more tubular appearance. It is
not possible to assign a species to a genus on the basis
of the external penis appearance. The internal struc-
tures must be examined. They differ radically from
subfamily to subfamily, and within a genus are subject
to extensive modifications for species recognition pur-
poses. A review of structure on a subfamily by subfam-
ily basis is required.
In the Charopinae from Australia, New Zealand,
New Caledonia, Lord Howe Island, South Africa, St.
Helena, and South America there is an amazing
amount of variation that will have to be dealt with
elsewhere. Comments here will be restricted to the
type genus, Charopa Albers, 1860, Phenacohelix Suter,
1892, and the Pacific Island taxa. Charopa coma (Gray,
1843) has the penis with only a modestly swollen ex-
terior (fig. 9b) and a very short basal stalk portion.
Internally (fig. 9d) it shows an apical verge (PV) with
near terminal slitlike pore, below this is a pocket
stimulator (PC), followed by a series of variable-sized
circular pilasters that occupy a quite low position. In
addition, the upper and middle walls of the penis
chamber have raised, soft pustules. Phenacohelix
pilula (Reeve, 1852) has a much elongated and more
slender penis (fig. lie), with shifted position of the
epiphallus (E) to below the insertion of penial retractor
muscle (PR). Internally (fig. lie) a small verge (PV)
with terminal pore lies above a series of corrugated
longitudinal pilasters and a vague swelling that I
interpret as a remnant of the pocket stimulator. The
verge and ridge correspond to the external bulge on the
penis section. The sculpture of the wall, position and
number of circular ridges, size of pocket stimulator,
and verge size differ between Phenacohelix and
Charopa, but the fact of a basic set of structures is the
important data.
In most of the Pacific Island Charopinae there is
simple variation in the relative sizes of the verge,
pocket or modified pocket stimulator, and the number
and prominence of the circular bands. The wall
sculpture found in the New Zealand taxa seems to be
absent in the Pacific Island taxa. The combination of
very small size (see table XV), limited material, and
preparation of illustrations at an early stage in the
study means that treatment of the variations in this
portion of the study is inadequate. Specific comments
on differences between geographically close species are
given in appropriate places in the systematic review.
Here I will mention only a few more obvious changes.
The verge varies from a small spade-shaped papilla
with open side (fig. 42e, S. peasei), to a conical projec-
tion with terminal pore (fig. 43e, S. auanaensis), or it is
reduced to a swelling attached to one wall (fig. 55e, S.
allecta), or a globular swelling (fig. 57c, S. com-
plementaria, fig. 75h, Ba humbugi), a protruding lobe
with laterally apical pore (fig. 79c, Tuimalila pilsbryi),
or the bulbous structure seen in Vatusila tongensis (fig.
84b).
The circular ridge can be split into several and low
(fig. 42d-e, Sinployea peasei), doubled and thick (fig.
43e, S. auanaensis), single and thin (figs. 51h, S. inter-
media; 57c, S. complementaria; 67j, S. euryomphala),
or very thick and partly altered into pads (figs. 55e, S.
allecta, 79c, Tuimalila pilsbryi).
The pocket stimulator is particularly variable and
with few exceptions has been inadequately illustrated
and interpreted. Comparison of the doughnut shape in
Figure 39h, split "U" in Figure 42d, raised globular
mass in Figure 53c, low "U" in Figure 53h, lobular
structure in Figure 55e, "U" ridge in Figure 67d,
widely open pocket in Figure 67j, greatly reduced and
altered structure in Figure 75h, and nearly closed
pocket in Figure 79d gives an idea of the great extent of
variation in this structure, but this is only an introduc-
tion to its complexity.
This study demonstrates that extensive variation
occurs in verge, circular ridges, and pocket stimulator
in the Pacific Island Charopinae, but analysis of the
patterns of variation is beyond the scope of this report.
Because Sinployea is evidently in a highly active stage
of speciation, study of this area might yield important
data, but is left for others.
A major modification of this pattern occurs in
Graeffedon (fig. 87c). The epiphallus (E) opens through
a simple pore that is surrounded by a huge circular
pilaster (PP). Immediately below this are three large
pilasters: an upper, free-tipped vergic stimulator, a
long and low semicircular ridge oriented longitudi-
nally in the penis chamber, and a broad, low ridge with
slightly free upper tip that occupies the lower half of
the penis. Coupled with the absence of the valve at the
vas deferens-epiphallus junction, these structures
mark Graeffedon as remote from the other Charopinae,
but it is better accommodated in this subfamily than
assigned to one of its own.
The Trukcharopinae (fig. 90c-d, g, i) have a simple
pore opening from the epiphallus (Russatus), a low
vergic papilla (Kubaryiellus), or even a tubular verge
(Palline). Various pilasters and a possibly highly mod-
ified pocket stimulator are found in some of the taxa.
These variations are discussed more fully on pp. 205-
207. Many of these represent additive structures com-
pared with the Charopinae, but basically they could be
derived from some of the Charopinae conditions in ex-
tralimital taxa (Solem, unpublished data).
The Semperdoninae (figs. lOOd, f, j, 102d, 103b, f)
have the epiphallus entering the penis through a nar-
row muscular collar, with high lamellar pilasters ex-
tending down and then usually coalescing into three,
high glandular pilasters. The whole penis is sur-
rounded by a muscular sheath. Exceptions to the basic
pilaster pattern are Himeroconcha rotula (Quadras &
Mollendorff, 1894), which lacks the initial radiating
pilasters although retaining the three basal ones, and
Ladronellum mariannarum (Quadras & Mollendorff,
30
SOLEM: ENDODONTOID LAND SNAILS
1894) in which the initial radiating pilasters coalesce
into a huge horseshoe-shaped pilaster (fig. 102d) that is
inflatable with fluids. The enclosure of both epiphallus
and penis by a muscle sheath is as great a change as is
the pilaster pattern. These make the Semperdoninae
the most isolated of the charopid subfamilies in terms
of penis structure.
In the few dissected Rotadiscinae (H. B. Baker,
1927; Solem, unpublished data) the epiphallus enters
through a pore, papilla, or short verge into a very
thick-walled chamber with longitudinal pilasters.
There may or may not be accessory organs associated
with the penis and/or atrium. This represents yet
another series of experiments in the Charopidae.
Finally, the situation in Discocharopa (fig. 34c) re-
quires comment. A short, tubular penis with lateral
entrance of the vas deferens has the penial retractor
muscle inserting apically. Apparently there are at
least two longitudinal pilasters inside the penis, but
the available material was not adequate to work out
the details (see p. 75). Although this departs from the
reported pattern for the Charopinae (see above), the
genus is temporarily placed in the Charopinae pending
revision of the extralimital groups. It probably will be
split into another subfamily unit, but available data
are inadequate to propose a more appropriate classifi-
cation.
The atrium, in all dissected material, is a simple,
short tube opening externally behind and above the
right rhinophore. It varies somewhat in length but
shows no really significant changes.
TERMINAL FEMALE GENITALIA The post-
uterine or free oviduct (UV), spermatheca (S), and va-
gina (V) in the Endodontidae are very slender tubes
without unusual structures. They show variation pri-
marily where the spermathecal shaft inserts penis,
atrium, or free oviduct (Solem, 1976b, pp. 83-84). In
the Charopidae the situation is quite different. Nor-
mally, the spermathecal base and at least parts of the
vagina are greatly enlarged and with complex internal
pilasters. The free oviduct, most of the vagina, and the
spermathecal shaft above the swelling will be quite
slender, with an ovate, expanded head of the sperm-
atheca buried in the albumen gland-prostate margin.
The shape and proportions shown by Sinployea avan-
aensis (fig. 43c) are typical. The shifted expanded
area in S. aunuuana (fig. 53b) or expanded free oviduct
(UV) in S. allecta (Cox, 1870) (fig. 55c), shortened va-
gina in S. inermis lakembana (fig. 67f), and generally
greater elongation in the Semperdoninae (figs. lOOb, e,
g, 102a, 103a, c-d) are the typical minor variations.
The nearest thing to a major variation is found in
Jokajdon (fig. 93bc) where the expanded area is re-
stricted to the base of the vagina and atrium, and the
normally expanded areas are slender in comparison. In
addition there is an accessory muscle attaching to the
penioviducal angle. These changes may correlate with
the very restricted shell aperture and large barriers of
Jokajdon.
Internally there appear to be two basic patterns of
sculpture of the upper vagina and lower spermathrca.
In the typical Charopinae, such as S. complementaria
(Mousson, 1865) (fig. 57f), S. allecta (Cox, 1870), and
Ba humbugi, there are weak longitudinal pilasters in
the vagina, the free oviduct (UV) has a constricting
pilaster with central pore (UVO), and the opening of
the spermatheca (S) has a central pore (SO) through a
circular pilaster with one edge a free flap. Because of
size problems and the early stage in the study at which
this area was examined, only the one has been illus-
trated. The second pattern is seen quite clearly in
Graeffedon (fig. 87c) where the opening to the free
oviduct is a simple pore and there are huge longitudi-
nal pilasters lining the walls of the spermatheca and
vagina. Essentially the same pattern is seen in
Palikirus, Palline, Semperdon uncatus, S. hepta-
ptychius (Quadras & Mollendorff, 1894), Ladronellum
mariannarum (Quadras & Mollendorff, 1894),
Himeroconcha lamlanensis, and H. fusca (Quadras &
Mollendorff, 1894), thus indicating that this general
type occurs not only in some of the Charopinae, but
also in the Trukcharopinae and Semperdoninae. A de-
tailed comparative study of this region would be --veil
worthwhile but is beyond the scope of this monograph.
PALLIAL COMPLEX
A discussion of the functional significance that the
closed and complete ureter found in most Charopidae
has in relation to water conservation as contrasted to
the incomplete posteriorly opening ureter of the En-
dodontidae was given previously (Solem, 1976b, pp.
84-87). Some of the New World Rotadiscinae show
only a partial closed ureter (H. B. Baker, 1927, pi. 16,
fig. 17, pi. 17, figs. 24, 30), and there are Australian
equivalents (Solem, unpublished data). All of the
Pacific Island Charopidae that have been dissected
show a complete ureter with the ureteric pore opening
next to the anus just inside the pneumostome. This
contrasts immediately with the Endodontidae where
the ureteric pore opens at the posterior part of the pal-
lial cavity near where the anterior margin of the rectal
kidney lobe touches the hindgut (Solem, 1976b, p. 85,
fig. 49c). Thus, even a glance at the anterior part of the
pallial cavity is sufficient to tell whether a Pacific Is-
land species belongs to the Endodontidae or
Charopidae.
A typical charopid configuration is seen in
Sinployea allecta allecta (Cox, 1870). Viewed from the
outside (fig. 55a), the bilobed nature of the kidney (K)
is obvious, with the longer, cigar-shaped rectal lobe
(lower in figure) definitely overlapping the hindgut
(HG). The shorter, irregularly shaped pericardial lobe
(upper in figure) partly overlies and is cupped partly
around the heart (H). The kidney base (left in figure)
stops short of the downward twist of the hindgut as
intestine and is abutted by the lobules of the digestive
gland. The primary ureter (KD) originates from near
the anterior margin of the pericardial lobe of the kid-
PATTERNS OF MORPHOLOGICAL VARIATION
31
ney, follows the upper margin of this lobe posteriorly,
lying partly on the pallial cavity roof and partly on the
kidney itself, reflexes abruptly and as the secondary
ureter follows the lower margin of the rectal kidney
lobe anteriorly to its termination, then lies next to the
hindgut until both disappear under the mantle collar
(MC). The principal pulmonary vein (HV) extends an-
teriorly from the heart along the pallial roof, but in the
smaller species shows no sign of branching, fading out
from visual observation well short of the mantle collar.
The bilobed kidney, or in a torn and extracted
specimen, the tip of the rectal lobe of the kidney, plus
the presence of the secondary ureter as a distinct tube
next to the hindgut, are sufficient to immediately iden-
tify a species as a member of the Charopidae or
Punctidae rather than the Endodontidae. In the latter
family (Solem, 1976b, p. 459, fig. 195a) there is at most
a short rectal lobe with the ureter ending in a ureteric
pore (KX) as it reaches the hindgut. In a few
Charopidae, such as Phenacohelix pilula (Reeve, 1852)
(fig. lla), the rectal arm is as short as in the Endodon-
tidae, but the clear presence of the secondary ureter
along the hindgut is sufficient for family separation.
Exceptions to this in some Australian taxa will be con-
sidered elsewhere.
When dissected out (fig. 55b) and viewed from an
inside view of the pallial cavity, the typical charopid
pallial region shows only minor additional features.
Remnants of a retractor muscle to a rather strongly
developed muscle come off the parietal-palatal margin
near the kidney base. A weaker version of the same
muscle can be seen in many species of Endodontidae
(Solem, 1976b, p. 459, fig. 195a). The heart in this view
clearly lies on top of the kidney, the rectal kidney lobe
extends partly under the hindgut, thus overlapping on
both sides of this tube, and both the hindgut and ureter
terminate just inside the mantle collar. Quite possibly
this area is involved in water resorption and is a pre-
cursor of the more elaborate structures seen in higher
aulacopods, such as an apparent bladder in Deroceras
reticulatum (Miiller, 1774) (Runham & Hunter, 1970,
pp. 77-79, figs. 32-33).
Modifications of this general pattern involve addi-
tion of new structures, changes in the relative size of
the kidney lobes, differences in the amount of space
between the two lobes, and compactional alterations
correlated with reduction in whorl counts. Data on at
least part of the pallial complex was recorded for 36 of
the 43 taxa for which at least some anatomical mate-
rial was available. Illustrations are presented for 20 of
these. The only additive structure seen in a Pacific Is-
land taxon is the extensive intrusion of mantle gland
tissue onto the pallial roof in Graeffedon graeffei
(Mousson, 1869) (fig. 87a, MG) and a slight extension
in Semperdon xyleborus. This phenomenon is far more
common in extralimital taxa, with the intrusion rang-
ing from a massive and sharply defined area in
Champa coma (Gray, 1843) (fig. 9a) to the short area in
Phenacohelix pilula (Reeve, 1852) (fig. lla) and the
elongated finger in Laoma leimonias (Gray, 1850) (fig.
25a). The function of this extension is unknown.
Although there is a modest amount of intraspecific
variation in the relative length of the two kidney lobes,
much of this seems to be caused by differential contrac-
tion and compaction when the animal withdraws into
the shell. Dissections made from deeply retracted
specimens tended to have the rectal kidney lobe un-
changed in shape and position, but pulled back further
relative to the heart and pericardial kidney lobe. The
latter would tend to be somewhat twisted and distorted
(for example see fig. 34d). I made no exact measure-
ments as to the relative lengths because the results
would not be strictly comparable. Table IX summarizes
the relative lengths in the Pacific Island species. Only
in Discocharopa aperta (Mollendorff, 1888) (fig. 34d) is
the rectal lobe somewhat reduced to as great an extent
as in the New Zealand Phenacohelix pilula (Reeve,
1852) (fig. lla). The general pattern is for the rectal
lobe of the kidney to be much longer than the pericar-
dial. This reaches its greatest extent in such taxa as
Graeffedon graeffei (Mousson, 1869) (fig. 87a) and
Semperdon heptaptychius (Quadras & Mollendorff,
1894) (fig. lOOa). In both of these taxa the pericardial
lobe is reduced to a small fraction of the length and
volume of the rectal.
The length relationship varies within a genus,
since in Sinployea there are nine species in which the
TABLE IX. - RELATIVE LENGTH OF KIDNEY LOBES
IN THE PACIFIC ISLAND CHAROPIDAE.
Subequal or
Equal
Sinployea tahitiensis
^. lamellicosta
j>. avanaensis
^. intermedia
S^. allecta allecta
ji. vicaria vicaria
^. kusaieana
j>. inermis inermis
J[. i_. lakemba
^. adposita
Ba humbugi
Tulmalila pilsbryi
Russatus nigrescens
Himeroconcha fusca
H. lamlanensis
Rectal Much
Longer
Sinployea modicella
^>. montana
S, neglecta
S. peasei
S^. aunuuana
^. clista
S_. complement aria
S^. ir_regulaj^is_
S^. eu r y ompha 1 a.
Graeffedon graeffei
Kubaryiellus kubaryi
Trukcharopa trukana
Palikirus cosmetus
Jokajdon calllzonus
Palline np_t era not era
P_. micramyla
Semperdon uncatus
j^. xyleborus
^. heptaptychius
S_. rotanus
Ladronellum raariannarum
Pericardial
Much Longer
Discocharopa aperta
32
SOLEM: ENDODONTOID LAND SNAILS
kidney lobes are equal or nearly equal in length and
nine in which the rectal is significantly longer. I could
detect no conchological correlations in size, shape, or
whorl count with this variation in kidney lobe length
concerning Sinployea. The other Charopinae have a
secondarily shortened and very fat kidney correlated
with whorl count reduction (Ba humbugi, fig. 75a),
subequal lobes in the gigantic Tuimalila pilsbryi (fig.
79a), or the very elongated rectal lobe in Graeffedon
graeffei (Mousson, 1869) (fig. 87a). In the Truk-
charopinae only Russatus nigrescens (Mollendorff,
1900) (fig. 90a), whose whorl count is reduced to a
mean of 3Vs + , has a shortened kidney, whereas in the
Semperdoninae only the large-sized Himeroconcha
have the kidney lobes equal or subequal in length.
In both the Trukcharopinae and the Semper-
doninae the arms of the ureter are tightly pressed to-
gether without any pallial roof tissue visible between
them. The pattern is somewhat unusual in Russatus
nigrescens (Mollendorff, 1900) (fig. 90a), with the
abrupt angling of both ureter arms at the anterior
margins of the kidney lobes. I interpret this as the
result of shortening and secondary thickening of the
kidney in relation to whorl and pallial cavity reduc-
tion. In the Charopinae, the same compaction is seen in
Ba humbugi (fig. 75a), with only slightly less abrupt
and extensive angling of the ureter. Of the examined
Sinployea, all taxa except S. allecta allecta (Cox, 1870)
(fig. 55a-b) and S. irregularis (Garrett, 1887) have at
least a narrow strip of pallial roof tissue visible be-
tween the ureter arms, as does Tuimalila pilsbryi (fig.
79a). Extralimital Charopinae are variable in this fea-
ture (see figs. 9e, lla).
Shortening of the pallial cavity is more common
than elongation. The latter seems to have occurred
only in Jokajdon callizonus (Mollendorff, 1900) (fig.
93a) where it extends % of a whorl apically. This prob-
ably correlates with the narrowed cross-section of the
body whorl in this species (fig. 92b). Major shortening
has occurred in Ba humbugi (fig. 75a) and Russatus
nigrescens (Mollendorff, 1900) (fig. 90a). Their respec-
tive mean whorl counts of 3% and 3Vs+ are the lowest
in the Pacific Island Charopidae, except for the rotadis-
cine Microcharopa mimula (mean whorl count 3V4-).
In both Ba and Russatus the width of the kidney is
almost equal to its length, and the arms of the ureter
are tightly compacted and overlap each other between
the kidney lobes. The process has been carried further
in Russatus, with V4 whorl length to the pallial cavity,
than inBa, where the length is still Vz whorl. The same
type of compaction has occurred in both genera, and
they form a contrast to the situation in the New
Guinea charopids Pilsbrycharopa and Paryphantopsis
(Solem, 1970a, p. 250, fig. 2a, f) in which kidney com-
paction resulted in progressive increase of the angle
between the arms of the ureter, and the kidney is ro-
tated away from the hindgut. Yet another pattern is
seen in the New Zealand flammulinid and Maori-
concha groups.
The pallial cavities in the Pacific Island Endodon-
tidae and Charopidae have radically different ureter
and kidney structures. Within the Pacific Island
Charopidae there is a relatively simple pattern of vari-
ation in relative lengths of the kidney lobes and the
extent to which lung roof tissue is visible between the
arms of the ureter.
DIGESTIVE SYSTEM
Allowing for the generally reduced whorl counts
(table X) in the Charopidae, the gross features of the
endodontid (Solem 1976b, pp. 372-373, figs. 163-164)
and charopid (figs. 9e, g, 75a) digestive tracts are the
same. The digestive glands (OG) are in contact above
the esophagus (BE), which continues past the pallial
cavity as a slender tube. In some taxa, such as the
larger Sinployea, Ladronellum mariannarum (Mollen-
dorff, 1900), and Kubaryiellus kubaryi (Mollendorff,
1900), the salivary glands are fused above the
esophagus, whereas in Ba humbugi, they do not even
touch posteriorly. I am not certain if there is any sys-
tematic significance to these changes. Intestinal loops
above the pallial cavity apex occupy about Vie of a
whorl, instead of the Vs whorl in the Endodontidae. The
charopid stomach expansion averages about % of a
whorl, with the digestive gland shorter and more com-
pact than in the Endodontidae. Except for these
changes associated with the reduced whorl count, the
gross features of the digestive tract are the same in the
two families.
Radular features are one of the best guides to
separating the Endodontidae, Charopidae, and Punc-
tidae on the Pacific Islands. The endodontid radula
(Solem, 1976b, pp. 88-94, figs. 51-54) has a tricuspid
central, several bicuspid laterals, and marginals that
have the ectocone fragmented and the endocone in-
creasing almost to the size of the mesocone. There are
very noticeable differences in the angling of the indi-
vidual teeth, but their small size and difficulties in
mounting and preparation have prevented full study of
their variation.
The punctid radula (fig. 12) presents a number of
unusual features. The central tooth (fig. 12a, upper)
has a long slender mesocone and two very slender,
much shorter ectocones. The laterals (fig. 12b-e) have
two slender cusps and three much shorter accessory
cusps, whereas the outermost lateromarginals (fig. 12f)
are broader and with the outer large cusp reduced in
size. The accessory cusps are at to below the limit of
optical microscope examination, depending upon the
quality of the equipment and illumination. As far as I
am aware, the only optical microscopist to detect and
illustrate these accessory cusps was H. B. Baker (1927,
pi. 16, fig. 11), also reprinted in Pilsbry (1948, p. 642,
fig. 349, d).
In addition to the obvious cusp edges and numbers
there are fundamental differences in the nature of the
cusps and basal plates in the two families. The en-
dodontid basal plate (Solem, 1976b, p. 89, fig. 52d) has
a typical interlocking relationship with the next pos-
FIG. 12. Radular teeth ofPunctum minutissimum (Lea). Cedar bog, Woodburn Road, 4 miles southwest of Urbana, Champaign County, Ohio.
E. Keferl! X-20-1969. FMNH 151102: a, central (upper) and 1st lateral (23,000x); b, early lateral teeth (10,800x); c, single midlateral tooth
(21,500x); d, late lateromarginal teeth (15,900x); e, low angle view of lateromarginal teeth (10,700x); f, outer lateromarginal tooth showing
cusp reduction (28,000x).
33
34
SOLEM: ENDODONTOID LAND SNAILS
terior tooth, is square to rectangular in shape (Solem,
1976b, p. 91, fig. 54b, e), and, at least for the early
lateral teeth, the cusps are sharply pointed and ele-
vated at a high angle (Solem, 1976b, p. 88, fig. 51a-b;
p. 89, fig., 52a-b, d). In the Punctidae (fig. 12c, e) the
basal plate is long and tapering, has no apparent inter-
locking relationships, and the cusps are bluntly
rounded and point almost directly forward. It is unfor-
tunate that these differences can be seen only with the
scanning electron microscope, a fact that reduces their
routine utility in identification and classification.
The Pacific Island Charopidae have a very stan-
dard pattern of structure (figs. 13-14). As pointed out
earlier (Solem, 1976b, p. 93), extralimital charopids
show a variety of structure, but review of these is out-
side the scope of this monograph. The tricuspid central
tooth (figs. 13b, 14a) is slightly narrower and shorter
than the adjacent laterals, which are tricuspid. When
viewed at a low angle, it is evident that the side cusps
of both central and laterals are raised above the eleva-
tion plane of the mesocone (fig. 14a), whereas in the
Endodontidae (Solem, 1976b, p. 88, fig. 51a) they are in
the same elevation plane. The outer laterals and early
marginals in the Charopidae (fig. 14b) do not have the
elevated side cusps. The marginals may retain the
tricuspid pattern (fig. 13c-f), or rarely the outer mar-
ginals may become multicuspid. Climo (1969a, figs.
31-34; 1970, figs. 11-14) gives a number of radular
transects for New Zealand charopids.
Because only a few species could be examined with
the SEM and their pattern of structure was quite
uniform, results from optical viewing mostly have been
omitted from the text in this volume.
The bicuspid laterals of the Endodontidae, tricus-
pid laterals of the Charopidae, and multicuspid
lateromarginals of the Punctidae found on Pacific Is-
lands thus present clearcut differences among the
families. The Austro-Zelandic charopids, however,
show a great variety of radular structures, so that the
endodontid-charopid distinction does not hold for that
area.
Jaw structure in the Charopidae was not studied
in detail because the pattern of partial plate fusion in
larger taxa paralleled the situation found in the En-
dodontidae (Solem, 1976b, p. 94).
FREE MUSCLE SYSTEM
As in the Endodontidae (Solem, 1976b, p. 94), all
dissected charopids had the right ommatophoral re-
tractor passing through the penioviducal angle and
joining the right rhinophoral retractor that passes out-
side the penioviducal angle posteriorly. Unlike the En-
dodontidae, in all examined Charopidae the penial re-
tractor muscle arises from the diaphragm and inserts
onto the penis or epiphallus. Only a few alterations in
the common pattern of unions were observed. Russatus
nigrescens (Mollendorff, 1900) has the tentacular re-
tractors fusing with the tail fan at the columellar
muscle rather than earlier as in most taxa. InJokajdon
callizonus (Mollendorff, 1900) a new muscle attaches at
the penioviducal angle, joining the tail fan much later.
With the elongated (% whorl) pallial cavity and nar-
rowed aperture with large barriers (fig. 92b) in this
species, the added muscle may play a major role in
successful body retraction.
In Sinployea complementaria (Mousson, 1865)
there is a muscle from the right tentacular retractor
that inserts on the apex of the free oviduct, whereas in
Tuimalila pilsbryi a muscle runs from the columellar
retractor to the apex of the free oviduct. It is possible
that a weaker version of this same muscle is present in
smaller species of Pacific Island Charopidae and was
overlooked in earlier phases of this study. It was not
possible to recheck all taxa for this feature. Extralimi-
tal taxa, such asStephanoda binneyana (Pfeiffer, 1847)
(fig. 31c), show a vaginal retractor muscle (VRM). It is
probable that a number of such experiments in added
muscles exist, but have not been observed.
NERVOUS SYSTEM
Wherever possible, the enervation of the penis
from the right cerebral ganglion was confirmed, but
because of difficulties in handling the small-sized
material, preservation in alcohol, and heavy covering
of connective tissue over the ganglia, no attempt at
working out the details of the nervous system was
made. Climo (1970, fig. 21B) illustrated the central
nervous system ofPhenacharopa novoseelandica (Pfeif-
fer, 1853).
EXTERNAL BODY FEATURES
In the Trukcharopinae and Semperdoninae, except
for Semperdon heptaptychius (Quadras & Mollendorff,
1894), Ladronellum mariannarum (Quadras & Mol-
lendorff, 1894), and Himeroconcha fusca (Quadras &
Mollendorff, 1894), the body color is yellow-white,
without darker markings. In the latter three species
there are gray to reddish gray markings on the neck,
ommatophores, and mantle collar. In the Charopinae,
all live-collected Cook, Society, and Samoan Island
species of Sinployea, except the Swains Island S. in-
termedia, have light to dark gray markings. All of the
Sinployea with gray markings have been taken in
semiarboreal situations. The purely terrestrial S.
kusaieana, S. euryomphala, S. inermis, S. adposita,
and S. irregularis (Garrett, 1887) have yellow-white
body color. Tuimalila pilsbryi also shows the darker
mantle and neck coloration. The dark body color ap-
pears to be a correlative of semiarboreal habitat.
Many of the arboreal New Zealand and Australian
charopid taxa have a strongly developed mucus appa-
ratus at the hind end of the foot. A typical "mucus
pore" or "caudal foss" (CF) is present in Phenacohelix
pilula (Reeve, 1852) (fig. lib), with the pore overhung
by a caudal horn (CH). This is effectively an intensifi-
cation of the point where the foot grooves unite above
the tail plus a mucus-secreting gland concentrating at
FIG. 13. Radular teeth of Tuimalila pilsbryi. Station T-22, 1,000ft. elevation, Eua, Tonga. FMNH 152378: a, near middle of radula at 280 x;
b, central tooth and 1st lateral on right side of radula at 2,820 x ; c, early laterals on right side of radula at 2,900 x ; d, transition from laterals to
marginals on left side of radula at 2,950x; e, middle marginals from left side at 2,875x; f, outermost marginals at 2,875x. All views nearly
vertical.
35
36
SOLEM: ENDODONTOID LAND SNAILS
FIG. 14. Radular teeth ofTuimalilapilsbryi. Station T-22, 1,000 ft. elevation, Eua, Tonga. FMNH 152378: a, 45 angle view of central (c) and
early laterals at 3,200 x showing pattern of cusp elevation; b, same view of transitional zone between laterals and marginals at 3,125x.
this point. This apparatus is absent in Charopa coma
(Gray, 1843) (fig. 9a) and all of the dissected Pacific
Island Charopidae (fig. 43a). The presence of this pore
is associated with arboreal snails (Solem, 1976b, pp.
105-106) and is not of major phyletic importance, as
was also confirmed by Pilsbry (1892a-b, 1893a-b) and
Climo (1969a, pp. 148-150).
As in the Endodontidae (Solem, 1976b, p. 94), the
mantle collar normally is without protrusions, the
gonopore is located below and slightly behind the right
ommatophore, and the slime network is rather weakly
defined. Neither family shows substantial external
modifications in the Pacific Island taxa.
PATTERNS OF VISCERAL HUMP REDUCTION
In the Endodontidae (Solem, 1976b, pp. 94-98) one
of the major repetitive changes is an increase in the
number of whorls and thus elongation of the visceral
hump. Not only do the Pacific Island Charopidae have
a much lower average whorl count than the Endodont-
idae (median means 4Vfe and 5V2 + , respectively), but
of the Charopidae only Semperdon kororensis (Bed-
dome, 1889) with 5Vs whorls and the Rarotongan
Sinployea planospira (Garrett, 1881) with 6% whorls
exceed a mean count of five whorls. The latter has not
been taken in this century, and of the former only
fragmentary extracted pallial collars and terminal
genitalia were available. Thus I can offer no data of
any changes associated with elongation of the visceral
hump.
Reduction in whorl count and thus in the total
length of the visceral hump is a common pattern in
charopids from many areas of the world. Such taxa as
the Juan Fernandez A mphidoxa (fig. 30a, c), New Zea-
land Flammulina, Maoriconcha, and Otoconcha
(Climo, 1971a), a few of the South African Trachycystis
(Connolly, 1939), and some New Caledonian taxa
(Solem, 1961) show varying degrees of visceral hump
reduction. This is carried furthest in the New Zealand
Otoconchinae and the semislug Ranfurlya. Discussion
of their changes is beyond the scope of this review, but
the general pattern is for zonal compaction of organs in
several systems (see Solem, 1966, for a discussion of
this in the Thailand Helicarionidae).
In the Pacific Island Charopidae whorl reduction
takes two forms simple decrease in mean whorl count
without any change in whorl profile (Microcharopa
mimula, Z l A-;Lagivala minusculus, 3%; Discocharopa
aperta, 3%; and Palikirus ponapicus, 3%) or reduction
in whorl count accompanied by a drastic increase in
cross-sectional areas of the body whorl (Russatus ni-
grescens, 3Vs + ',Ba humbugi, 3%).
Either the former taxa have not been dissected, or,
in the case of Discocharopa, there are no close relatives
known with which anatomical comparisons can be
made. Thus, comments here must be restricted to the
latter situation, whorl count reduction combined with
whorl cross-section increment. The degree of whorl
changes can be judged by comparing whorl increment
rates and whorl profiles in Kubaryiellus and Russatus
(fig. 89a-b, d-e) and then in Sinployea irregularis
(Garrett, 1887) (fig. 65d-e) and Ba humbugi (fig.
74a-b). Anatomically, the typically trukcharopinine
half whorl pallial region of Kubaryiellus (fig. 90e) can
easily be altered to the nearly square kidney (K) and
one quarter whorl pallial cavity of Russatus (fig. 90a) by
shortening and widening of the kidney, ventral flexing
PATTERNS OF MORPHOLOGICAL VARIATION
37
of the intestinal loop, and a slight ventral movement of
the anterial part of the pericardial kidney lobe over the
heart (H) and principal pulmonary vein (HV). Compar-
ing the gross genitalia (fig. 90b, f), the proportionately
shortened free oviduct (UV), spermatheca (S), prostate
(DG), and uterus (UT) ofRussatus are obvious. There is
no detectable change in the penis (P) and vagina (V).
The pallial region of Sinployea irregularis (Garrett,
1887) was not illustrated, but it has the typical generic
pattern of extending % of a whorl apically, there is no
lung roof space visible between the arms of the ureter,
and the rectal lobe is distinctly longer than the
pericardial. In Ba humbugi (fig. 75a) the anterior half
of the half whorl pallial cavity has undergone shorten-
ing and change, with the anterior margins of the kid-
ney flared laterally, but the posterior portion of the
kidney and the intestinal loops are essentially unmod-
ified when compared with the degree of change seen in
Russatus (fig. 90a). Contrasting the genitalia of S. ir-
regularis (fig. 67a, drawn from a deeply retracted
specimen) and So (fig. 75b-e, h) shows a rather drastic
folding of the prostate and uterus plus altered insertion
of the penial retractor muscle in the latter taxon. Ba
also shows shortening of the vagina and spermatheca.
Thus, changes in the visceral hump length of
Pacific Island Charopidae involve selective shorten-
ings in portions of the organ systems lying in the pal-
lial region. Unless there is clear shortening of the neck
region (area between ommatophores and pallial collar
edge when animal is crawling), the prostate-uterine
area and spermathecal-free oviduct sections are more
apt to be involved than the penis-vagina, and the kid-
ney area, than the gas exchange surfaces of the pallial
roof. More extended comments on these changes are
postponed pending completion of studies on Australian
and New Zealand taxa.
SUMMARY OF ANATOMICAL VARIATION
The Pacific Island Charopidae show a number of
minor variations in the terminal genitalia and kidney
configurations. The former involve species recognition
factors, the latter are of uncertain significance except
in cases of obvious elongation or shortening of the pal-
lial cavity. These terminal genitalia patterns are not
continuously variable, but fall into rather discrete
general patterns that indicate multiple colonizations of
the island areas and are used in part to recognize sub-
family units.
The major and minor anatomical differences be-
tween the Pacific Island Charopidae and Endodontidae
were summarized by Solem (1976b, pp. 97-98, tables
LVIII-LIX). Consideration of the complexities in the
Australian, New Zealand, and New Caledonian taxa
must be deferred. There are no clear unitary anatomi-
cal trends within the Pacific Island Charopidae com-
pared with those outlined for the Endodontidae (Solem,
1976b, pp. 98-99), although the addition of structures
among the various charopid subfamilies is striking.
A major correlative of the "looser" whorl coiling
pattern (pp. 41-43) in the Charopidae compared with
the Endodontidae is that the cross-sectional area of the
body whorl in particular is greater in the former fam-
ily. This has important implications on the size of the
anterior body and its contained organs. The wider area
in the Charopidae permits widening of the terminal
genitalia. The much fatter penis and vagina-free
oviduct-spermathecal union area in the Charopidae is
possible directly because of this extra space provided
by the looser coiling. Whereas in the Endodontidae
these organs all are thin tubes, in most Charopidae
they are thick and with complex internal structures.
The only Pacific Island charopid known to have
clearly narrowed structures in this area is Jokajdon
callizonus (Mollendorff, 1900) (fig. 93a-b), whose dras-
tically narrowed shell aperture and large apertural
barriers (fig. 92a) closely approach the typical en-
dodontid condition. Even though species of Palline
(figs. 94b, e, 95d) have large apertural barriers and
somewhat narrowed apertures, the thick terminal
genitalia (fig. 93f, h) contrasts with that of Jokajdon
and is in the typical charopid pattern.
It is quite probable that the variety of genital
structures seen in the Charopidae as opposed to the
Endodontidae are in large part the result of simple
space availability for experimentation.
CHAROPID-ENDODONTIDCONCHOLOGICAL
COMPARISONS
Despite almost complete overlap with regard to
most conchological characteristics and their close simi-
larity when viewed with the naked eye, there are a few
clear differences between the Endodontidae and
Charopidae in meristic and structural features. Data
in Tables I-III and X and Figures 15-23 summarize
both similarities and differences in some basic para-
meters. Figures 15-22 were prepared several years ago
and omit data on the Lau Archipelago endodontids,
Priceconcha tuuuthaensis Solem (1973d) and Thauma-
todon spirrhymatum Solem (1973d). Their inclusion
would not have changed the results significantly. In
addition, four taxa of endodontids were not seen or
measured. These two exceptions account for the
difference between the 185 taxa listed by Solem
(1976b, p. 9, table IV) and the 179 listed as measured in
this tabular comparison.
For ease in visual comparison, in Figures 15-22
the actual numbers of species for each graph unit have
been converted into percentages of the total measured
within that family so that the graphs will be directly
comparable. Otherwise the difference between 179 en-
dodontids and 95 charopids would make visual com-
parisons difficult.
Table X shows the median mean value and total
range of mean values for the species measured within
each family. The distinctly larger size and higher
whorl count of the Endodontidae is evident, whereas
their near identity in H/D ratios, D/U ratios, and mean
rib counts is surprising. The greater ribs/mm, in the
Charopidae directly correlate with their smaller size.
TABLE X. - CONCHOLOGICAL COMPARISONS OF ENDODONTIDAE AND CHAROPIDAE
Number of species
level taxa measured
Median mean height (range)
Median mean diameter (range)
Median mean whorl count
(range)
Median mean H/D ratio
(range)
Median mean D/U ratio
(range)
Median mean ribs on body
whorl (range)
Median mean ribs/mm.
Taxa with ribs reduced
PACIFIC ISLANDS.
Endodontidae
179
1.48(0.92-7.26)
3.77(1.7-12.3)
5 1/2+
(3 5/8-8 1/8)
0.531
(0.344-0.789)
3.94
(1.68-closed)
91.0(19-202)
9.2(1.4-40)
39(21.8%)
Charopidae
95
1.51(0.48-3.69)
2.76(1.07-7.52)
4 1/8-
(3 1/8-6 5/8)
0.523
(0.365-0.801)
3.94
(2.03-closed)
91.2(19-225)
11.3(1.7-37)
10(10.5%)
The greater degree of rib reduction in the Endodon-
tidae and the reduction in percentage of taxa with
apertural barriers in the Charopidae also indicate
major differences.
More detailed comments can be made from the
data in Figures 15-23. Mean height distribution (fig.
15) shows a slight Endodontidae offset for most species,
then an extended high-spired portion that far sur-
Charopidae
Endodontidae
175 275 375
Mean Height
4.75 575 over 65
38
FIG. 15. Mean shell height distribution in the Endodontidae and
Charopidae.
passes the maximum height recorded for the Char-
opidae. The three highest spired Charopidae are
Lauopa mbalauuana, Tuimalila infundibulus (Hom-
bron & Jacquinot, 1841), and T. pilsbryi. All three have
average to only slightly increased whorl counts, and
only the latter has a high SP/BWW ratio (mean 2.23).
Their large height is the result of size increase alone,
rather than change in shell form. In contrast, the
high-shelled Endodontidae mostly are species with
umbilical brood chambers and increased whorl counts,
plus the few Nesodiscus and pre-brood chamber En-
dodonta (Solem, 1976b, pp. 27-30).
Mean diameter distribution (fig. 16) shows a
greater similarity between the two families, both hav-
ing a significant right extension of the frequency
curve. The sharper peak for the Charopidae relates to
CHAROPID-ENDODONTID CONCHOLOGICAL COMPARISONS
39
30
25 -
20 -
Charopidae
Endodontidae
1.75
2.75 3.75
475
Mean Diameter
5-75 6.75 7.75 over
8.0
FIG. 16. Mean shell diameter distribution in the Endodontidae
and Charopidae.
the speciose genus Sinployea, which accounts for more
than half of the total taxa reviewed. The secondary
peak around 4.75 mm. results from some of the large
Rarotongan and Samoan Sinployea, plus Himero-
concha, Russatus, and large Semperdon from Micro-
nesia. Both families have a few taxa that exhibit
gigantism compared with the average species.
Mean H/D ratio distribution (fig. 17) differs only
because of the brood chamber taxa in the Endodontidae
(Solem, 1976b, p. 29, fig. 19), producing a slight bulge
30
25
20
~
I
o 15
-
I
-Charopidae
Endodontidae
0.325
0.425
0.525 0-625
Mean H/D Ratio
0.725
0.825
FIG. 17. Mean height/diameter ratio distribution in the En-
dodontidae and Charopidae.
in the higher ratios for that family. The only very
high-spired charopids are the nearly scalariform Ba
humbugi (fig. 74b), Ladronellum mariannarum (Qua-
dras & Mollendorff, 1894) (fig. lOlb), and Semperdon
kororensis (Beddome, 1889) (fig. 98b). Only Ladronel-
lum has a normally open umbilicus (fig. lOlc), whereas
the others have a closed umbilicus that normally in-
creases the H/D ratio dramatically (Solem, 1976b, p.
25, fig. 15).
The most dramatic difference is shown by the
mean whorl count distribution (fig. 18). The only
charopids to exceed a mean whorl count of five whorls
Charopidae
Endodontidae
( I
10
3 1 '.
4'i
5'', 6'*
Mean Whorls
FIG. 18. Mean whorl count distribution in the Endodontidae and
Charopidae.
are the Micronesian Semperdon kororensis (Beddome,
1889) (fig. 98a) with 5V& average, and the Rarotongan
Sinployea planospira (Garrett, 1881) (fig. 46d) with 6%
whorl average. Although there is considerable overlap
in mean whorl counts of five whorls or less, the drama-
tic increase in higher mean whorl counts in the En-
dodontidae is quite clear. A fair portion of this increase
to the right correlates with brood chamber formation
(Solem, 1976b, p. 29, fig. 20).
Means of D/U ratio for the Endodontidae in Figure
19 exclude the brood chamber taxa entirely, hence the
two families show very similar patterns. Because egg
laying in the umbilicus is a family characteristic in the
40
SOLEM: ENDODONTOID LAND SNAILS
n
25-
20-
Charopidae
Endodonlidae
225 325 4.25 525 6.25 7- 11- 21- crack cloud
10 20 50
X D/U Ratio
FIG. 19. Mean diameter/umbilical width ratio distribution in the
Endodontidae and Charopidae.
Endodontidae (Solem, 1976b, pp. 100-101), extreme
narrowing or closure of the umbilicus should be a rarer
event in the Endodontidae than in the Charopidae.
Surprisingly enough, it is not. The only Charopidae
with a closed or nearly closed umbilicus are Semperdon
kororensis (Beddome, 1889) (fig. 98c) from Palau,
Sinployea clista (fig. 51c) from Samoa, and Be humbugi
(fig. 74c) from Viti Levu, Fiji. Sinployea clausa (fig.
5 If) from Samoa has an extremely narrowed um-
bilicus. Most other species have the umbilicus much
more widely opened, and only in the Lau Archipelago
Sinployea adposita (Mousson, 1870) (fig. 69f) andRuss-
atus nigrescens (Mollendorff, 1900) (fig. 89f) from
Ponape is the umbilicus very narrow. In the Endodon-
tidae (Solem, 1976b, pp. 26, 491-492) there are 10 taxa
with closed and 10 with barely perforate umbilici. Most
of these are found on Rapa and Mangareva, areas far
from the main areas of distribution and possibly from
the natural occurrence of potential egg predators that
occupy the leaf litter. Hence, the unexpected high
occurrence of closed or barely perforate umbilici in the
Endodontidae is a geographic phenomenon.
The degree of spire protrusion, as measured by the
SP/BWW ratio, is quite different for the two families
(fig. 20). The Endodontidae are in general much higher
spired. This relates primarily to the taxa reaching the
Nesodiscus and brood chamber levels of organization.
The most elevated Charopidae are Ba humbugi from
Fiji (fig. 74b), Ladronellum mariannarum (Quadras &
Mollendorff, 1894) from Guam (fig. lOlb), Sinployea
angularis from Fiji (fig. 64e), and Vatusila nayauana
from Fiji (fig. 82e). Because so few Pacific Island
Charopidae have elevated spires, the type of analysis
done for the Endodontidae (Solem, 1976b, p. 25, fig. 15)
in which spire protrusion was correlated with varia-
Cfiaropidae -.-
Endodontidae .
depressed Hal 0.05 0.15 0.25 035 04 055 065 075 085 0.95 overlO
Spire height/body whorl width ratio
FIG. 20. Mean spire height/body whorl width ratio distribution
in the Endodontidae and Charopidae.
tions in shell height, diameter, H/D ratio, and D/U
ratio is not presented since the differences were so
slight. Comparatively few Charopidae have a flat or
clearly depressed spire. Only Roimontis tolotomensis
has a clearly sunken spire.
The striking similarity in median mean rib counts
on the body whorl (table X) and the very similar distri-
bution of mean rib counts in the two families (fig. 21)
was not anticipated. It does suggest that the hypoth-
esized function of the ribbing to reduce adherence of
particles to the shell surface (Solem, 1976b, p. 50) may
be correct. Rib reduction in the Endodontidae is
primarily size correlated. Species with a mean diame-
ter of 4.75 mm. or more frequently show a marked de-
gree of rib reduction (Solem, 1976b, pp. 46-50, tables
Cliaiofxd*
Endodontidae
10 -
190 over reduce)
200
70 110 ISO
Ribs on Body Whorl
FIG. 21. Mean ribs on body whorl distribution in the Endodon
tidae and Charopidae.
CHAROPID-ENDODONTID CONCHOLOGICAL COMPARISONS
41
XVII- XIX). There are comparatively few taxa in the
Charopidae that show major rib reduction. The 10 are
not phyletically correlated and show reduction in
different degrees and ways. In Himeroconcha rotula
(Quadras & Mollendorff, 1894) and H. lamlanensis
from Guam, plus Sinployea rudis (Garrett, 1872) and
S. harveyensis (Garrett, 1872) from Rarotonga, the
sculpture is reduced to irregularity on the body whorl,
with the major ribs becoming too crowded and too ir-
regular to count well before the lip. In Sinployea ir-
regularis (Garrett, 1887) and Ba humbugi from Fiji
plus Russatus nigrescens (Mollendorff, 1900) from
Ponape, the postapical sculpture is highly irregular,
and occasionally a rib with a high lamellar extension
appears. In Sinployea recursa from Fiji and
Himeroconcha quadrasi (Mollendorff, 1894) and H.
fusca (Quadras & Mollendorff, 1894) from Guam, the
sculpture is reduced in prominence on both the spire
and body whorl and rapidly becomes indistinguishable
from growth lines after an initial portion where the
ribs are large enough to be counted. The four
Himeroconcha do show a pattern of size-associated re-
duction in rib prominence, since the two smaller
species have more prominent spire sculpture (fig.
104a-f) than do the two larger taxa (fig. 105a-f). There
is no data available concerning the ecology of
Himeroconcha, so that the reasons for this pattern of
sculpture reduction are unknown. Russatus, Ba, and
Sinployea irregularis (Garrett, 1887) are known to be
terrestrial in habitat, and Garrett (1872, pp. 227-228)
reported that both S. rudis (Garrett, 1872) and S. har-
veyensis (Garrett, 1872) were collected under rotting
wood. Only dead examples of the Lau Archipelago
Sinployea recursa are known, although I have specu-
lated that it could be an arboreal species.
The overall pattern of rib reduction in the
Charopidae is less clearly size linked than in the En-
dodontidae. Three of the 10 species, S. recursa, S. ir-
regularis (Garrett, 1887), and Ba humbugi are within
0.11 mm. of the median mean diameter, the other
seven (38.9%) are among the 18 over 3.75 mm. in
diameter, and three (37.5%) are among the eight
species that average more than 4.75 mm. in diameter.
They are Russatus nigrescens (Mollendorff, 1900),
Himeroconcha fusca (Quadras & Mollendorff, 1894),
and H. quadrasi (Mollendorff, 1894). In the Endodon-
tidae, 50% of the taxa averaging over 4.76 mm. in
diameter have reduced sculpture (Solem, 1976b, p. 47,
table XVIII).
The variation in mean ribs/mm, on the body whorl
(fig. 22) is equivalent in the two families, once allow-
ance is made for the smaller mean size of the Charo-
pidae. As in the Endodontidae (Solem, 1976b, pp.
44-45, tables XV- XVI), the smaller the mean diame-
ter, in general the more numerous are the radial ribs
(see table X).
In summary, basic shell size, shape, and radial
sculpture spacing and frequency in both families are
more similar than dissimilar, once allowance is made
Charopidae
[ndodonlidae
J I
H 15
Ribs/mm
I I
over reduced
22
FIG. 22. Mean ribs/mm, on the body whorl distribution in the
Endodontidae and Charopidae.
for the changes in the Endodontidae caused by the for-
mation of a brood chamber. The Endodontidae are
larger and with a higher whorl count than the
Charopidae, but in basic form and character of the ra-
dial sculpture they are very similar. This is not sur-
prising because both are basically found in litter,
under stones, and in or under rotting wood. A much
higher number of Charopidae than Endodontidae has
been taken in semiarboreal to arboreal situations.
In addition to gross size, shape, and sculpture
comparisons, an important question is whether there is
a different pattern of growth between the two families.
With the complications introduced by variations in
spire protrusion, body whorl descension, umbilical
width, whorl counts, and sculptural protrusions, it is
difficult to find a simple index of whorl increment pat-
tern that will at least indicate the pattern of whorl
width increase. Figure 23 attempts this through use of
a crude measure of size increment plotted against
whorl count of the figured individual. To try and
minimize difficulties in interpretation, brood chamber
taxa and the very high-spired members ofAaadonta in
the Endodontidae plus the high-spired Ba humbugi in
the Charopidae have been omitted. Inclusion of these
would have added data points to the lower right of the
figure. Because the increase in shell diameter is less
when the spire is elevated, the results would have been
artificially skewed toward the right.
The basic data for Figure 23 were taken from the
top views of shells as published in Solem (1976b) and
this report. Because top views were not published for
species of Anceyodonta, nearly all Cookeconcha, Opan-
ara, Rhysoconcha, Ruatara, Orangia, Taipidon,
Planudonta, Rikitea, and Nesophila, this is only a par-
tial sampling of the Endodontidae. In contrast, nearly
all of the Charopidae have been figured in top view. On
each illustration, two measurements were made: the
width of the first whorl from suture to suture and then
the diameter of the entire shell. In addition, the
42
SOLEM: ENDODONTOID LAND SNAILS
27-
26
25-
24
23
22
21
20
19
18
I 16
Endodontidae
Charopidae
D
D D
O D
D DD
DO D D
DD Q
OOD 6
D
5
2
D
D D D
3
n
D DO OOD dS
13
12
11
D
DQ D rm
4
D I I OT
D DO DQ
D DD
D D
10
D
an
Whorl count
FIG. 23. Patterns of whorl width increment in selected Endodontidae and Charopidae. Brood chamber, very high spired, and unillustrated
top views of species in the Endodontidae are omitted, as is the high-spired Ba humbugi in the Charopidae. Unusual taxa are: (1) Minidonta
manuaensis Solem, 1976; (2) Mautodontha aoraiensis Solem, 1976; (3) Australdonta pseudplanulata Solem, 1976; (4) A. pharcata Solem, 1976; (5)
Cookeconcha stellulus (Gould, 1844); (6) Thaumatodon laddi Solem, 1976; (7) Zyzzyxdonta alata Solem, 1976; (8) Aaadonta kinlochi Solem, 1976;
(9)Sinployea canalis (Garrett, 1872); (10) S . planospira (Garrett, 1881); (11) Russatus nigrescens (Mollendorff, 1900); (12) Semperdon kororensis
(Beddome, 1889); and (13) Minidonta micraconica Solem, 1976.
number of whorls in the illustrated specimen was re-
corded. The width of the first whorl was divided by the
total diameter to give a percentage of the total diame-
ter taken up by the initial nuclear whorl. This is, of
course, partly inaccurate, because the suture-to-suture
distance will be less than the periphery-to-periphery
distance. The error will be approximately the same for
each species, so that the degree of error will be in the
same direction. Use of a percentage index of total
diameter plotted against the actual whorl count gives
an approximate indication of basic whorl width incre-
ments.
It is evident that members of the Endodontidae
have a tighter pattern of coiling at the same whorl
count than do the Charopidae. The higher the percent-
age of the first nuclear whorl width, the slower is the
rate of whorl width increment for succeeding whorls.
There is, in general, a rather clear separation between
the two family units. The exceptions from the general
pattern in the Charopidae are a few species with un-
usual coiling patterns: (number 9 on fig. 23) Sinployea
canalis (Garrett, 1872) (fig. 49a-c), which has a later-
ally compressed whorl profile and increased whorl
count; (number 10) S. planospira (Garrett, 1881) (fig.
CHAROPID-ENDODONTID CONCHOLOGICAL COMPARISONS
43
46d-h), which is laterally compressed, has increased
whorl count, and very tight coiling; and (number 12)
Semperdon kororensis (Beddome, 1889) (fig. 98a-c),
with its raised spire, lateral compression, and closed
umbilicus. At the other extreme, (number 11) Russatus
nigrescens (Mollendorff, 1900) (fig. 89d-f), which has a
flat spire, reduced whorl count, and reduced sculpture,
is offset to the left considerably. The unusual Endodon-
tidae include (number 1) Minidonta manuaensis Solem
(1976b, p. 131, fig. 62a) and (number 13) Minidonta
micraconica Solem (1976b, p. 138, fig. 65a-c) with re-
duced whorl counts and abnormally flat-spired taxa in
normally more elevated genera such as (number 2)
Mautodontha aoraiensis Solem (1976b, p. 160, fig.
74d-f), (number 3) Australdonta pseudplanulata Solem
(1976b, p. 295, fig. 127d-f), and (number 4) A.pharcata
Solem (1976b, p. 313, fig. 137a-c). Taxa whose diame-
ter was increased by greatly enlarged radial ribs in-
clude (number 5) Cookeconcha stellulus (Gould, 1844)
(Solem, 1976b, p. 218, fig. 93a-c) and (number 7) Zyz-
zyxdonta alata Solem (1976b, p. 466, fig. 198a-c).
Aaadonta kinlochi Solem (1976b, p. 486, fig. 208a-c,
number 8) is a flat-spired species in a genus that is
normally highly elevated, whereas (number 6)
Thaumatodon laddi Solem (1976b, p. 452, fig. 193d-f)
is a flat-spired species with slightly protruded
periphery. Thus, all exceptions are readily explained
as representing special situations.
Correlated with the more rapid whorl width in-
crement in the Charopidae is a greater increase in
cross-sectional area of the whorl profile. Without sec-
tioning shells and measuring the areas of the whorls,
quantification of this difference is not feasible. The fact
of the greater cross-sectional area and concomitant in-
crease in linear wall distance at any given point un-
doubtedly has had major effects on the placement of
pallial organs in relation to each other and their
length. It also has permitted thickening of genital or-
gans over the endodontid condition. Discussion of these
changes was dealt with under the patterns of anatomi-
cal variation.
On the submicroscopic level, there are three major
differences between the Endodontidae and Charopidae
found on the Pacific Islands. The apical sculpture of the
shell, the method of forming the postapical shell
sculpture, and the way in which the apertural barriers
are formed and armed with microdenticles are
different. These differences have been reviewed in part
elsewhere (Solem, 1969d). Only a brief summary is in-
cluded here.
In the Endodontidae, the apical sculpture consists
of prominent radial ribs, with or without microriblets
in between, plus very fine spiral cords that are best
termed "squiggly" (Solem, 1976b, pp. 35-41, figs.
25-31). In one genus, Aaadonta Solem (1976b, pp.
38-39, figs. 28-29), the major radial sculpture has
been lost, only the microradials are left on the postapi-
cal whorls, and only the squiggly spiral cords are left
on the apex. In the Pacific Island Charopidae, the basic
apical sculpture is of strong spiral cords, typically as
seen in Sinployea modicella (Ferussac, 1840) (fig.
la-c). They can be reduced in prominence and in-
creased in number, as in Sinployea peasei (fig. 2a-b), or
combined with a secondary sculpture of low radial
swellings, as in Tuimalila pilsbryi (fig. 2c). The only
exceptions on the Pacific Islands concern the rotadis-
cine genus Microcharopa, in which the spiral apical
cords are broken up into short, twisted segments (fig.
4a-e), and the widespread Discocharopa (fig. 5), in
which both apical and postapical sculpture consists of
fine radial ribs and there is no trace of spiral sculpture.
The situation becomes much more complicated in the
charopid taxa of Australia and New Zealand where
there have been many experiments in shell sculpture
(Solem, unpublished data). However, in relation to the
Pacific Island taxa, the sharp division into taxa with
spiral cords (Charopidae) and taxa with radial ribs
(Endodontidae) holds with a few secondary exceptions.
Under optical magnification, Aaadonta would be con-
fused with the charopid condition and Tuimalila with
the endodontid condition, but SEM studies show that
these apparent exceptions are secondary modifications.
Except for obvious periostracal setae and exten-
sions in such taxa as Cookeconcha decussatulus (Pease,
1866) (Solem, 1976b, p. 36, fig. 26a-c; p. 40, fig. 30b),
the postapical sculpture in the Endodontidae appar-
ently is formed by a thin template of periostracum,
with most of the sculpture thickness consisting of un-
derlying calcium layers. This holds even for the fine
apical features (Solem, 1976b, p. 35, fig. 25d). In con-
trast, the microsculpture and much of the major rib
projection in the Charopidae consists entirely of perio-
stracal materials. Frequently the only calcareous
sculptural element will be a swelling underneath each
major radial rib. An example of this is seen in
Sinployea uicaria vicaria (Mousson, 1871) (fig. 59). The
same pattern of structure seems to hold true for at least
many of the New Zealand and Australian Charopidae.
This makes the similarity in gross sculpture effect be-
tween the two families even more remarkable.
An initial review of the apertural barrier
differences between the Endodontidae and Charopidae
was given in Solem (1973b). The following summary is
taken from that paper, Solem (1976b, pp. 52-72), and
pp. 15-23. In the Endodontidae, the barriers show a
uniform pattern of structure, growth, and microden-
ticulation. The parietal and columellar walls will have
weak pustulations, as is typical of many taxa in a vari-
ety of families (Solem, 1972c), but the characteristic
feature on the barrier tops is a series of additive tri-
angular microdenticulations (Solem, 1973b, figs. 7-13,
23-24; Solem, 1976b, p. 55, fig. 39d-e; p. 64, figs. 40,
41a-c). It apparently does not matter whether the ex-
panded upper edge of the barrier is continuous or
broken up into a series of expanded beads as in
Thaumatodon, Aaadonta, and Zyzzyxdonta (Solem,
1976b, p. 457, fig. 194b-e; p. 466, fig. 198b; p. 475, fig.
203b, e). The pattern of minute triangular denticles
44
SOLEM: ENDODONTOID LAND SNAILS
that face toward the aperture and are additions to the
surface of the barrier, rather than an outgrowth from
it, holds. The only known exceptions are in certain of
the very large species of Cookeconcha and Endodonta,
where the denticulations on the upper edge of the bar-
riers are blunt-tipped (Solem, 1976b, p. 66, fig. 42d-f)
although retaining their triangular shape on the sides
of the barriers, and in other genera such asNesodiscus,
where the barriers are greatly reduced in both size and
number. These barriers may be secondarily without
denticulations. I have thus concluded that the barriers
seen in members of the Endodontidae had a common
origin and that reduction or loss of barriers is a secon-
dary phenomenon in that family.
In the Charopidae (pp. 15-17), in contrast, three of
the four major groups with barrier-equipped genera
found on the Pacific Islands show barriers that differ in
size, position, growth, and microdenticulations (Solem,
1973b, pp. 303-304, figs. 4-6, 14-22). The latter basi-
cally are crystals growing out of the barrier surface
and are blunt tipped. It is thus concluded that aper-
tural barriers have evolved independently in each of
these lineages, despite the fact that SEM observations
could not be made on the Smp/ojea-derivative genera.
Their barriers show such obvious differences from the
other three groups (figs. 80-83, 85) that I have no
doubt of their independent origin. More detailed dis-
cussion of the barrier structure and variation has been
given in the Patterns of Morphological Variation (pp.
15-23).
In the Endodontidae the great majority of the
species have barriers on the palatal wall, with a
bladelike shape and gradual anterior descension. In
the Charopidae only four species have bladelike
palatal barriers, and 13 have purely crescent-shaped
barriers.
The frequency of barrier occurrence is lower in the
Charopidae than in the Endodontidae. In the latter,
including taxa not seen but illustrated in older litera-
ture, 184 out of 185 (99.5%) have parietal barriers and
163 (88.1%) have palatal barriers. In the Charopidae
32 (33.7%) of 95 taxa have parietals, and 27 (28.4%)
have palatal barriers. Thus the families differ signifi-
cantly not only in whether or not barriers are present
in the aperture but also in how they are formed, the
normal shape of the palatals, and the nature of the
microdenticulations on the barrier edges.
Despite great optical similarity in size, color,
shape, presence of apertural barriers in many species,
and presence of fine radial ribbing in at least the
smaller taxa, differences in whorl count (fig. 18),
growth pattern of whorl width (fig. 23), apical
sculpture, method of formation for the postapical
sculpture, and details of the apertural barrier
structure and microdenticulations serve to distinguish
the two families on conchological grounds.
HABITAT RANGE AND EXTINCTION
In contrast with the Endodontidae (Solem, 1976b,
pp. 100-101), the Charopidae have been able to exploit
the semiarboreal and arboreal habitats as well as re-
maining well represented in the ground stratum. Their
closed secondary ureter has been hypothesized (Solem,
1976b, p. 100) as the reason the charopids have been
able to become successfully semiarboreal. A second fac-
tor in their success is the apparent lack of umbilical
egg deposition by the charopids. Of all the Pacific Is-
land charopid specimens studied, only one example of
Tuimalila infundibulus (Hombron & Jacquinot, 1841)
had a snail egg capsule inside the umbilicus, whereas
this is the routine pattern in the Endodontidae. Of ex-
tralimital taxa, the New Zealand charopids, Fectola
marsupialis Powell, 1941, and Aeschrodomus worleyi
Powell, 1928, have the umbilical egg-laying pattern
(see Climo, 1969a, pp. 218-219; 1970, p. 306).
On such drastically ecologically altered islands as
Viti Levu, Tahiti, and Moorea, specimens ofSinployea
and Ba have been collected from lowland vegetational
remnants in the 1960s and 1970s. They have not, how-
ever, been found where the ground stratum has been
severely and regularly disturbed by chickens and/or
pigs. In parts of Samoa and on the Lau Archipelago of
Fiji the charopids still are common and relatively
abundant. Only on Rarotonga is there clear evidence of
major charopid extinctions within a century. Two sepa-
rate collecting efforts on Rarotonga in the mid-1960s
failed to find nine out of 10 Sinployea species collected
in the 1860s and 1870s and five out of six endodontids
reported from the same island. All of the missing taxa
were described as ground stratum inland species. The
one collected endodontid occupies the storm line coral
boulder habitat, and the one previously described
Sinployea is at least occasionally semiarboreal.
The actual egg-laying habits of the Pacific Island
Charopidae are unknown, but their nondependence
upon the umbilicus for egg-laying probably has effec-
tively lowered ant predation, which apparently was a
major factor in the mass extinction of the Pacific Island
Endodontidae. The semiarboreal to arboreal habitat
range extension of the Charopidae also has lessened
human impact through introductions, because arboreal
snails of the Pacific Islands, in general, have been able
to adapt to plantations of bananas and even coffee,
whereas strictly ground-dwelling taxa are absent from
these human-introduced environments.
The charopids are abundant in Australia, South
Africa, New Zealand, New Caledonia, and parts of
South America all areas in which ants are common. I
thus expect that the ability of the Pacific Island
charopids to survive despite the introduction of preda-
cious ants is a carryover from conditions in their areas
of greater abundance rather than a new adaptation.
Since ants first appeared in the Cretaceous fossil rec-
ord (Wilson et al., 1967), and have an extensive fossil
record of living tribes and genera from the Oligocene
and Miocene, their coexistence with the basic charopid
stock seems highly probable.
The charopids thus occupy a wider range of
habitats than do the Endodontidae and persist in fair
variety despite the great ecological alterations of the
past century on the Pacific Islands.
45
PHYLOGENY AND CLASSIFICATION
Reviews of the hypothesized early evolution and
progressive trends among land snails have been given
previously (Solem, 1974; 1976b, pp. 102-104; 1978b).
The position of the Endodontidae as the most gen-
eralized extant group of the Sigmurethra, the absence
of any family unit from which the endodontid anatomi-
cal structures can be derived, and the positioning of
both the Charopidae and Endodontidae within the
Arionacea complex have been discussed earlier and
will not be rejustified here.
The Charopidae are more advanced than the En-
dodontidae in having a closed secondary ureter, fused
prostate-uterine tubes with a common lumen, a re-
duced number of lobes in the ovotestis, and in transfer-
ring sperm in a packet (see Ba humbugl, fig. 75f-g),
which correlates with the general presence of an
epiphallus. Charopids are more complex than the En-
dodontidae in having great elaboration of terminal
genital structures and in the variety of their radular
structures. Recent dissections of some undescribed
Western Australian taxa have revealed genera that
are partly transitional between the Endodontidae and
Charopidae. These will be described and discussed
elsewhere.
It is premature to discuss in detail the probable
phylogenetic relationships among the endodontoid
families. After synoptic coverage of the Austro-
Zelandic taxa, such a discussion will be profitable. At
this time I choose to end the discussion of interfamily
phylogeny by emphasizing that the Charopidae are
more advanced than and derivable from the Endodont-
idae. The phylogenetic position of the Punctidae in re-
lation to the other families is still uncertain.
46
FAMILY CLASSIFICATION OF THE ENDODONTOIDS
A list of subfamily and family names available for
members of this complex was given in Solem (1976b, p.
105). I recognize five families and group the assignable
names as below. The name Patulinae Tryon (1866, p.
243) is discarded (see Solem, 1976b, p. 105). The af-
finities of the Indian Thysanotinae Godwin-Austen,
1907, are still unknown. Available literature informa-
tion on their anatomy is not adequate, and I have not
been able to obtain preserved materials for dissection. I
have not dissected one of Iredale's family units, the
Hedleyoconchidae, but on the basis of shell structures
have no hesitation in lumping it with the Charopidae
pending further study. Definitions and discussion of
the Helicodiscidae (see Solem, 1975) and Discidae
(Solem, in preparation A) are not included here, since
both families are extralimital to this study. Definitions
of both the Punctidae and Charopidae are included
below in the systematic review; the Endodontidae were
defined by Solem (1976b, p. 121).
The proposed synonymization of family level units,
in chronological order, is:
Family Punctidae (Morse, 1864, p. 27)
+ Laominae Suter (1913, p. 732)
+ Paralaomidae Iredale (1941a, p. 263)
Family Charopidae Mutton (1884b, p. 199)
+ Phenacohelicidae Suter (1892a, p. 270)
+ Otoconchinae Cockerel! (1893, pp. 188, 205)
+ Flammulinidae Crosse (1894, p. 210)
+ Rotadiscinae H. B. Baker (1927, pp. 226, 230)
+ Amphidoxinae Thiele (1931, p. 575)
+ Dipnelicidae Iredale (1937b, pp. 22-23)
+ Hedleyoconchidae Iredale (1942, pp. 34-35)
+ Pseudocharopidae Iredale (1944, p. 312)
Family Endodontidae Pilsbry, 1895 (Pilsbry, 1893-1895, p.
xxviii)
Family Helicodiscidae Pilsbry, 1927 (in Baker, 1927, pp. 226,
230)
+ Stenopylinae Thiele (1931, p. 569)
Family Discidae Thiele ( 1931, p. 578)
+ Goniodiscinae Wagner (1927, p. 305)
Although I do not accept strict nomenclatural
priority for names of families and higher level taxa, the
above usage comes close to following this precept. I
reject both Patulinae Tryon (1866) and Goniodiscinae
(Wagner, 1927); the former name because Patula
mostly has been associated with members of the land
snail family Oreohelicidae, a totally unrelated group,
until finally assigned as a synonym of Discus.
Goniodiscus Fitzinger, 1833, is now ranked as a sub-
genus or synonym of Discus (see discussion by Forcart,
1957), and I consider it inappropriate to use as a
family-level name.
The morphological gaps between these families
are of the same order of magnitude or greater than the
gaps separating recognized families of the Limacacea
(see Solem, 1976b, p. 107), although the gaps between
the family units certainly are not equal in size.
Discussion of subfamily and generic units is de-
ferred to the family discussions of the Punctidae and
Charopidae.
47
PHYLOGENETIC PROCEDURES
I am a pragmatic phylogeneticist in the sense of
Mayr and have used neither Hennigian cladistics nor
phenetic manipulations in this study. The criteria used
are those cited by Solem (1976b, p. 108) and the three-
tiered approach to evolutionary change outlined in
Solem (1978b). These are suited to the peculiarities of
molluscan shell growth, with the ontogenetic develop-
ment from embryo on visible in old age, the water-
dependent ecology of land snails, and their use of ter-
minal genitalia for species recognition.
The absence of many fixed reference points within
the snail's body and the difficulties of making precise
and meaningful measurements on the shell combine to
make quantification of many features impossible and
thus use of the more sophisticated mathematical ma-
nipulations of data matrices impractical. Similarly, in
the absence of equivalently detailed analyses of most
other land snail families and with the uncertainty as to
the overall phylogeny of either characters or taxa, use
of cladistic methodology is premature.
48
GENERIC CLASSIFICATION
The same type of "pigeon-hole" generic units cited
for the Endodontidae (Solem, 1976b, pp. 118-119)
existed for the Pacific Island Charopidae. If there were
apertural barriers present, the species was referred to
the form genus Endodonta or to one of its sections, and
if there were no such barriers present, it was placed in
the New Zealand genus Charopa. Although Pilsbry
(1893-1895, p. 21) had pointed out the artificiality of
this division, no significant changes were introduced
subsequently. Reference of a new species to Flam-
mulina (Russatus nigrescens) by Mollendorff (1900) or
attempts to use other described genera than Charopa
(Solem, 1959a, 1960) did not alter the essential absence
of any meaningful classification on the generic level.
Of the 21 genera in the Punctidae and Charopidae
reviewed below, only two, Punctum and Discocharopa,
have available names. The other 19 are named here.
The same criteria for generic separation are used here
in the Charopidae as were used in classifying the En-
dodontidae into genera (see Solem, 1976b, pp. 119-
120). Thus the generic concepts are directly compar-
able and the minimum generic gap equally applied.
The difference in frequency of monotypic genera in the
two families and presence of one particularly speciose
genus in the Charopidae was summarized by Solem
(1976b, p. 120). This is a real difference in evolutionary
patterns and not an artifact of classification.
49
SYSTEMATIC REVIEW
Because the two families are so disparate in the
number of species reviewed and the complexities of
charopid classification require considerable reference
to extralimital taxa, the discussion of charopid
phylogeny and classification is subtended to the family
review.
Following are a formal list of the taxa reviewed
and geographic keys to the genera. These sections fol-
low the format used in Solem (1976b) as do the species
and generic accounts.
LIST OF THE TAXA
Family Punctidae Morse ( 1864, p. 27)
Genus Punctum Morse, 1864
Punctum sp. Society Islands: Tahiti, Mt. Aorai
Punctum polynesicum, new species Austral Islands: Tubuai,
Raivavae
Family Charopidae Hutton (1884b, p. 199)
Subfamily Rotadiscinae H. B. Baker (1927, pp. 226, 228)
Genus Microcharopa, new genus
Microcharopa rnimula, new species Fiji Islands: Viti Levu; Lau Ar-
chipelago (Munia, Mothe, Wangava, Nayau, Namuka, Yangasa
Levu)
Subfamily Charopinae Hutton ( 1884b, p. 199)
Genus Discocharopa Iredale, 1913
Discocharopa aperta (Mollendorff, 1888) Philippines, Indonesia,
New Guinea, Western Australia, Northern Territory, Queensland,
Bismarck Archipelago, New Hebrides, Fiji Islands, Samoan Is-
lands, Austral Islands (Rurutu), Society Islands (Borabora)
Genus Sinployea, new genus
Sinployea modicella (Ferussac, 1840) Society Islands: Moorea
Sinployea tahitiensis, new species Society Islands: Tahiti
Sinployea lame/licosta (Garrett, 1884) Society Islands: Tahiti
Sinployea montana, new species Society Islands: Tahiti
Sinployea neglecta, new species Society Islands: Huahine
Sinployea sp. Society Islands: Borabora
Sinployea atiensis (Pease, 1870) Cook Islands: Atiu, Aitutaki
Sinployea andrewi, new species Cook Islands: Mangaia
Sinployea peasei, new species Cook Islands: Rarotonga
Sinployea avanaensis, new species Cook Islands: Rarotonga
Sinployea proximo (Garrett, 1872) Cook Islands: Rarotonga
Sinployea planospira (Garrett, 1881) Cook Islands: Rarotonga
Sinployea rudis (Garrett, 1872) Cook Islands: Rarotonga
Sinployea decorticata (Garrett, 1872) Cook Islands: Rarotonga
Sinployea harveyensis (Garrett, 1872) Cook Islands: Rarotonga
Sinployea youngi (Garrett, 1872) Cook Islands: Rarotonga
Sinployea canalis (Garrett, 1872) Cook Islands: Rarotonga
Sinployea otareae (Garrett, 1872) Cook Islands: Rarotonga
Sinployea tenuicostata (Garrett, 1872) Cook Islands: Rarotonga
Sinployea clausa, new species Samoan Islands: Manu'a Group
(Tau)
Sinployea clista, new species Samoan Islands: Tutuila, Upolu
Sinployea aunuuana, new species Samoan Islands: -Aunuu off
Tutuila, possibly Manu'a Group (Tau)
Sinployea allecta allecta (Cox, 1870) Samoan Islands: Upolu,
Savai'i
Sinployea allecta tauensis, new subspecies Samoan Islands: Manu'a
Group (Tau)
Sinployea complementaria (Mousson, 1865) Samoan Islands: Upolu
Sinployea intermedia, new species Swains Island
Sinployea vicaria vicaria (Mousson, 1871) Hoorn Islands: Futuna;
Tonga Islands: Vava'u, Tongatapu, Eua, Ha'apai Group
Sinployea vicaria paucicosta, new subspecies Tonga Islands:
Vava'u
Sinployea rotumana (Smith, 1897) Rotuma
Sinployea ellicensis ellicensis, new species and subspecies Ellice Is-
lands: Funafuti
Sinployea ellicensis nukulaelaeana, new subspecies Ellice Islands:
Nukulaelae
Sinployea pseudovicaria, new species Ellice Islands: Vaitupu
Sinployea kiisaieana, new species Caroline Islands: Kusaie
Sinployea sp. Mariana Islands: Saipan (possibly introduced)
Sinployea angularis, new species Fiji Islands: Lau Archipelago
(Namuka)
Sinployea recursa, new species Fiji Islands: Lau Archipelago
(Wangava)
Sinployea princei (Liardet, 1876) Fiji Islands: Taveuni, Kandavu
Sinployea inermis inermis (Mousson, 1870) Fiji Islands: Lau Ar-
chipelago (Vanua Mbalavu, Mango, Kimbombo)
Sinployea inermis meridionalis, new subspecies Fiji Islands: Lau
Archipelago (Yangasa Levu, Navutu-i-Loma, Aiwa)
Sinployea inermis lakembana, new subspecies Fiji Islands: Lau Ar-
chipelago (Lakemba)
Sinployea lauensis, new species Fiji Islands: Lau Archipelago
(Nayau, Yangasa Levu, Navutu-i-Loma, Wangava)
Sinployea navutuensis, new species Fiji Islands: Lau Archipelago
(Navutu-i-Loma, ? Oneata)
Sinployea adposita (Mousson, 1870) Fiji Islands: Lau Archipelago
(Oneata, Mothe, Munia, Lakemba, Aiwa, Karoni, Nayau, Yangasa
Levu)
Sinployea irregularis (Garrett, 1887) Fiji Islands: Viti Levu
Sinployea godeffroyana, new species Fiji Islands: Viti Levu
Sinployea euryomphala (Solem, 1959) New Hebrides: Espiritu
Santo, Maewo, Gaua, Vanua Lava
Sinployea solornonensis (Clapp, 1923) Solomon Islands: Ugi Island
off San Cristobal
Sinployea kuntzi (Solem, 1960) Solomon Islands: Florida Island off
Tulagi
Sinployea nissani (Dell, 1955) Solomon Islands: Nissan (Green Is-
land), north of Bougainville
Sinployea novopommerana (Rensch, 1937) Bismarck Archipelago:
New Britain
Sinployea descendens (Rensch, 1937) Bismarck Archipelago: New
Britain
Genus Ba, new genus
Ba humbugi, new species Fiji Islands: Viti Levu
Genus Maafu, new genus
Maafu thaumasius, new species Fiji Islands: Lau Archipelago
(Nayau)
50
SYSTEMATIC REVIEW
51
Genus Lauopa, new genus
Lauopo mbalavuana, new species Fiji Islands: Lau Archipelago
(Vanua Mbalavu)
Genus Tuimalila, new genus
Tuimalila infundibulus (Hombron & Jacquinot, 1841) Tonga Is-
lands: Vava'u
Tuimalila pilsbryi, new species Tonga Islands: Eua
Genus Lagivala, new genus
Lagivala davidi (Ladd, 1968) Ellice Islands: Funafuti (fossil)
Lagivala vivus, new species Fiji Islands: Viti Levu
Lagivala minusculus, new species Fiji Islands: Viti Levu
Lagivala macroglyphis (Rensch, 1937) Bismarck Archipelago: New
Britain
Lagivala microglyphis (Rensch, 1937) Bismarck Archipelago: New
Britain
Lagivala demani (Tapparone-Canefri, 1883) Indonesia (Timor,
Ambon, Aru Islands); West Irian (Misool)
Genus Vatusila, new genus
Vatusila kondoi, new species Fiji Islands: Lau Archipelago (Nayau)
Vatusila nayauana, new species Fiji Islands: Lau Archipelago
(Nayau)
Vatusila eniwetokensis (Ladd, 1958) Marshall Islands: Eniwetok
(fossil)
Vatusila tongensis, new species Tonga Islands: Eua
Vatusila vaitupuensis, new species Ellice Islands: Vaitupu
Vatusila niueana, new species Niue Island
Genus Graeffedon, new genus
Graeffedon graeffei (Mousson, 1869) Samoan Islands: Upolu
Graeffedon savaiiensis, new species Samoan Islands: Savai'i
Graeffedon pricei, new species Tonga Islands: Tongatapu
Subfamily Trukcharopinae, new subfamily
Genus Trukcharopa, new genus
Trukcharopa trukana, new species Caroline Islands: Truk,
Lukunor
Genus Kubaryiellus, new genus
Kubaryiellus kubaryi (Mollendorff, 1900) Caroline Islands: Ponape
Genus Russatus, new genus
Russatus nigrescens (Mollendorff, 1900) Caroline Islands: Ponape
Genus Roimontis, new genus
Roimontis tolotomensis, new species Caroline Islands: Ponape
Genus Palikirus, new genus
Palikirus cosmetus, new species Caroline Islands: Ponape
Palikirus ponapicus (Mollendorff, 1900) Caroline Islands: Ponape
Genus Jokajdon, new genus
Jokajdon tumidulus (Mollendorff, 1900) Caroline Islands: Ponape
Jokajdon callizonus (Mollendorff, 1900) Caroline Islands: Ponape
Genus Palline, new genus
Palline notera notera, new species and subspecies Palau Islands:
Peleliu
Palline notera palauana, new subspecies Palau Islands: Koror
Palline notera gianda, new subspecies Palau Islands: Babelthuap
Palline biakensis, new species West Irian: Biak
Palline micramyla, new species Caroline Islands: Ponape
Subfamily Semperdoninae, new subfamily
Genus Semperdon, new genus
Semperdon uncatus, new species Palau Islands: Angaur, Aulup-
tagel, Koror
Semperdon xyleborus, new species Palau Islands: Angaur, Koror,
Ngemelis, Peleliu
Semperdon kororensis (Beddome, 1889) Palau Islands: Koror
Semperdon rotanus, new species Mariana Islands: Rota, northern
tip of Guam
Semperdon heptaptychius (Quadras & Mollendorff, 1894) Mariana
Islands: Guam
Genus Ladronellum, new genus
Ladronellum mariannarum (Quadras & Mollendorff, 1894)
Mariana Islands: Guam
Genus Himeroconcha, new genus
Himeroconcha lamlanentiis, new species Mariana Islands: Guam
Himeroconcha rotula (Quadras & Mollendorff, 1894) Mariana Is-
lands: Guam
Himeroconcha quadrasi (Mollendorff, 1894) Mariana Islands:
Guam
Himeroconcha fusca (Quadras & Mollendorff, 1894) Mariana Is-
lands: Guam
Incertae sedis
Helix filiola Ferussac, 1840 Tonga Islands
Helix oceanica Le Guillou, 1842 Society Islands: Tahiti
Helix minutialis Deshayes, 1851 Society Islands: Tahiti
Helix multispirata Hombron & Jacquinot, 1852 Tonga Islands:
Vava'u
Helix rotula Hombron & Jacquinot, 1852 (not Lowe, 1831) Gambier
Islands: Mangareva
Pithys verecunda Pease, 1870 Marquesas
GEOGRAPHIC KEYS TO THE GENERA
The following artificial keys supplement those
presented for the Endodontidae in Solem (1976b, pp.
124-126). They are designed to enable identification of
adult shells. Endodontid taxa are cross-referenced
below but usually are not keyed down to the generic
level in order to save space and printing costs. Because
there are no charopids or punctids known from the
Mangareva, Marquesas, Rapa, or Tuamotu Islands,
these areas are not included in this set of keys; nor are
keys presented for extralimital areas. Additions to this
set of keys are the Caroline, Mariana, and main Fijian
Islands. For islands not included in the main groups,
reference to the list of "odd island" taxa at the end of
the keys may help.
AUSTRAL ISLANDS
1. Apical whorls with radial ribs 2
Apical whorls with spiral cords
Punctum polynesicum (p. 58)
2. Shell minute, diameter less than 1.8 mm.; at most 1 deeply
recessed parietal barrier
Discocharopa aperta (Mollendorff, 1888) (p. 76)
Shell normally much more than 1.8 mm. in diameter; usually
several apertural barriers
Endodontidae (Solem, 1976b, p. 125).
CAROLINE ISLANDS
1. Aperture without barriers 2
Aperture with 1 or more barriers 6
2. Whorls increasing rapidly in width (fig. 89d); mean D/U ratio
about 10; mean whorl count about 3 l /e
Russatus nigrescens (Mollendorff, 1900) (p. 215)
Whorls increasing less rapidly in width (fig. 89a); mean D/U
ratio 3.40-4.00; mean whorl count usually more than 3'/2. ...3
52
SOLEM: ENDODONTOID LAND SNAILS
3. TrukorKusaie 4
Ponape 5
4. Kusaie; spire elevated (fig. 63a); mean rib count on body whorl
about 91 Sinployea kusaieana (p. 143)
Truk; spire barely elevated (fig. 63d); mean rib count on body
whorl about 150 Trukcharopa trukana (p. 208)
5. Spire fiat or sunken; ribs on body whorl about 132; mean diame-
ter about 3.75 mm
Kubaryiellus kubaryl (Mbllendorff, 1900) (p. 212)
Spire slightly elevated; ribs on body whorl about 46; mean
diameter about 2.2 mm
Palikirus ponapicus (Mollendorff, 1900) (p. 221)
6. No columellar or palatal barriers 7
Palatal and/or columellar barriers present 8
7. Only 1 parietal barrier; spire elevated; 25 large ribs on body
whorl Palikirus cosmetus (p. 219)
8. Parietal barriers 2; spire sunken; about 60 low ribs on body
whorl Jtoimontis tolotomensis (p. 217)
Parietal and palatal barriers many and large 9
Barriers consist of 1 low parietal, 2 low palatals
Palling micramyla (p. 234)
9. Spire more elevated; ribs larger; body whorl rounded (fig.
92e) Jokajdon tumidulus (Mollendorff, 1900) (p. 223)
Spire barely elevated; ribs smaller; body whorl laterally com-
pressed behind aperture (fig. 92b)
Jokajdon callizonus (Mollendorff, 1900) (p. 226)
COOK ISLANDS
1. Apical whorls with radial ribs; at least 1 apertural barrier
Endodontidae (Solem, 1976b, p. 125)
Apical whorls with spiral cords; no apertural barriers
Sinployea, key on p. 98
ELLICE ISLANDS
1. Apical whorls with spiral cords 2
Apical whorls with radial ribs
Endodontidae, Thaumatodon decemplicata (Mousson, 1873)
(Solem, 1976b, p. 451)
2. Aperture with barriers 3
Aperture without barriers Sinployea, see p. 134
3. Vaitupu; living; 3 small barriers
Vatusila vaitupuensis (p. 196)
Funafuti; fossil; 6 large barriers
Lagivala davidi (Ladd, 1968) (p. 184)
FIJI, MAIN ISLANDS
1. Apical whorls with spiral cords 2
Apical whorls with radial ribs
Discocharopa aperta (Mollendorff, 1888) (p. 76)
2. Aperture without barriers 3
Aperture with barriers 5
3. Diameter less than 1.3 mm Microcharopa mimula (p. 71)
Diameter more than 2.0 mm 4
4. Spire greatly elevated (fig. 74b); umbilicus closed
Ba humbugi (p. 172)
Spire nearly flat or slightly elevated; umbilicus open
Sinployea, see key on p. 147
5. Parietal barriers 3; D/U ratio about 2.80; ribs on body whorl
about 62 Lagivala uiuus (p. 185)
Parietal barrier 1; D/U ratio about 5.25; ribs on body whorl
about 120 Lagivala minusculus (p. 185)
HAWAIIAN ISLANDS
Apical whorls with radial ribs, or, in larger species, smooth
Endodontidae (Solem, 1976b, p. 125)
Apical whorls with spiral cords; shell minute
Punctum horneri (Ancey, 1904) (p. 57)
LAU ARCHIPELAGO, FIJI
1. Apical whorls with radial ribs 2
Apical whorls with spiral cords 3
2. Several apertural barriers; diameter more than 2.5 mm
Endodontidae (Solem, 1976b, p. 125)
One or no apertural barriers; diameter less than 2 mm
Discocharopa aperta (Mollendorff, 1888) (p. 76)
3. Aperture without barriers 4
Aperture with 1 or more barriers 6
4. Diameter more than 2.0 mm 5
Diameter less than 1.3 mm Microcharopa mimula (p. 71)
5. Periphery protruded into a threadlike keel (fig. 76b); ribs on
body whorl about 44 and very large
Maafu thaumasius (p. 176)
Periphery at most angulated (fig. 68e); ribs reduced in size and
usually more numerous Sinployea, see key on p. 147
6. Diameter less than 2.0 mm 7
Diameter more than 7.0 mm.; 1 parietal barrier
Lauopa mbalavuana (p. 177)
7. Palatal barriers 3; parietal barrier bifid
Vatusila kondoi (p. 192)
Palatal barrier 1; parietal barrier simple
Vatusila nayauana (p. 194)
MARIANA ISLANDS
1. Aperture without barriers 2
Aperture with barriers 3
2. Shell small, diameter less than 2.5 mm.; Saipan
Sinployea sp. (p. 145)
Shell large, 3.8-7.2 mm.; Guam Jfimeroconcha (p. 259)
3. Aperture with 3 very small barriers (fig. lOlb)
Ladronellum mariannarum (Quadras & Mbllendorff, 1894)
(p. 255)
Aperture with 6 to many barriers (fig. 99b, d-e)
Semperdon rotanus and S. heptaptychius (Quadras & Mollen-
dorff, 1894) (pp. 245, 247)
MARSHALL ISLANDS
1. Apical whorls with spiral cords
Vatusila eniwetokensis (Ladd, 1958) (p. 195)
2. Apical whorls with radial ribs
Endodontidae (Solem, 1976b, p. 125)
PALAU ISLANDS
1. Postapical whorls with microradial riblets only, no major radial
ribs Endodontidae, Aaadonta (Solem, 1976b, p. 473)
Postapical whorls with major ribs and microsculpture 2
2. Shell diameter less than 2.6 mm.; mean whorl count less than
4V* 3
Shell diameter more than 2.7 mm.; mean whorl count more
than4V2 5
3. Palatal barriers 4, low ridges (figs. 94e, 95a); Koror and Babel-
thuap 4
Palatal barriers 5, high blades (fig. 94b); Peleliu
Palline notera notera (p. 229)
4. Palatal barriers 2; Koror Palline notera palauana (p. 231)
Palatal barriers 3; Babelthuap. ..Palline notera gianda (p. 231)
5. Palatal barriers many (fig. 98b, e) 6
Palatal barriers 0-1 (fig. 97b) Semperdon uncatus (p. 238)
6. Mean diameter about 4.6 mm.; umbilicus a very narrow
slit Semperdon kororensis (Beddome, 1889) (p. 244)
Mean diameter 2.9 mm.; umbilicus widely open
Semperdon xyleborus (p. 24P
SYSTEMATIC REVIEW
53
TABLE XI. - SPECIES NUMBERS OF
ENDODONTOID LAND SNAILS FROM SELECTED GEOGRAPHIC AREAS
PUNCTIDAE
CHAROPIDAE
Polynesia, Fiji
Hawaii 1
Known
1
1
Melanesia
New Guinea
Indonesia
New Caledonia'*
Lord Howe Id .
Norfolk Id.
Kermadec Is. 8
Australia
Tasmania
Victoria
N. S. Wales
Queensland
S. Australia
N. Territory
7
2
80
14
9
3
4
1
6
St. Helena
Neotropica and
Juan Fernandez
New Known
29
16
11
9
49
19
8
7
147
38
18
28
14
2
3
10
114
11
55(?)
2
TOTALS
646
Solem (1976b, this monograph, in preparation)
Solem (1959a, 1960, 1962, 1963, this monograph)
3 Solem (1958b, I970a)
4 Solem (1961)
Collecting by L. Price and P. Bouchet
Iredale (1944) and subsequent collecting
Iredale (1945) and subsequent collecting
8 Iredale (1913)
9 Powell (1976)
Iredale (1937a, 1937c, 1939)
U Burch (1976)
Collecting by A. Solem
New
ENDODONTIDAE
Known New
143
31
11
50(7, 5 ' 12
??
14
3
290
293
Connolly (1939); Solem (1970c)
Solem (1978a)
Various papers and Odhner (1922)
Pilsbry (1948) and various papers
F. Clirao, personal communication
SAMOAN ISLANDS
1. Apical whorls with radial ribs 2
Apical whorls with spiral cords 3
2. Aperture without palatal barriers; umbilicus very wide
Discocharopa aperta (Mollendorff, 1888) (p. 76)
Aperture with palatal barriers; umbilicus rather narrow
Endodontidae (Solem, 1976b, p. 126)
3. Aperture with prominent barriers 4
Aperture without barriers Sinployea, see key on p. 119
4. Shell diameter about 3.3 mm.; whorls 4Vi; Savai'i
Graeffedon savaiiensis (p. 202)
Shell diameter over 5.0 mm.; whorls about 5; Upolu
Graeffedon graeffei (Mousson, 1869) (p. 200)
SOCIETY ISLANDS
1. Apical whorls with radial ribs 2
Apical whorls with spiral cords 3
2. Shell diameter less than 2.0 mm.; at most 1 deeply recessed
parietal barrier
Discocharopa aperta (Mollendorff, 1888) (p. 76)
Shell diameter over 2.25 mm.; usually several parietal barriers
or shell diameter over 3.0 mm
Endodontidae (Solem, 1976b, p. 126)
3. Shell minute, diameter less than 1.5 mm....Punctum sp. (p. 57)
Shell larger, diameter over 2.4 mm Sinployea, see p. 86
TONGA
1. Apical whorls with spiral cords predominating, at most low ra-
dial swellings 2
Apical whorls with narrow radial ribs
Endodontidae (Solem, 1976b, p. 126)
2. Aperture without barriers 3
Aperture with prominent barriers 5
3. Diameter less than 3.5 mm Sinployea, see key on p. 134
Diameter more than 5.0 mm 4
4. Eua; mean diameter about 6.7 mm. ..Tuimalila pilsbryi (p. 179)
Vavau; mean diameter about 5.7 mm
Tuimalila infundibulus (Hombron & Jacquinot, 1841) (p. 181)
5. Palatal wall bearing a broad ridge (fig. 83b); Eua
Vatusila tongensis (p. 196)
Palatal wall with 3 barriers (fig. 88e); Tongatapu
Graeffedon pricei (p. 205)
A few species are known from islands that do not
fall into the major groups. These species are:
Futuna Sinployea vicaria vicaria (Mousson, 1871) (p. 134)
Niue Vatusila niueana (p. 198)
Rotuma Sinployea rotumana (Smith, 1897) (p. 138)
Swains Sinployea intermedia (p. 131)
It must be emphasized that the above keys are
adequate for identification of adult specimens in which
the sculpture and apertural barriers can be observed.
They are less adequate for juveniles and quite in-
adequate for very worn examples. It still will be possi-
ble to collect new species in areas such as the Caroline,
Palau, Mariana, Lau, and main Fijian Islands that
may or may not key out to a correct genus.
FAMILY PUNCTIDAE
Generally minute to small endodontoids, primitively with spiral
apical sculpture and a combination of macro- and microradial post-
apical sculpture that is periostracal in origin. Sculpture altered in
larger taxa. Whorl counts and shape highly variable, particularly in
New Zealand taxa. Umbilicus open, regularly decoiling to nearly
closed. Color monochrome to highly tessellated. A few taxa develop
apertural barriers, but most lack barriers. Foot of animal undivided.
Prominant pedal and suprapedal grooves unite above tail without
forming a caudal foss. Radula with small tricuspid central.
54
SOLEM: ENDODONTOID LAND SNAILS
lateromarginals bicuspid with three minute accessory cusps, major
cusps becoming reduced in prominence on outer teeth, but not tend-
ing to split into additional cusps. Basal plates long and slender, cusps
tending to point directly forward. Jaw of separated, square plates,
partly fused in larger species. Pallial complex with a bilobed kidney,
lobes generally equal in length, primary ureter originating at apex of
pericardia! kidney lobe, reflexing posteriorly as secondary ureter,
opening at ureteric pore alongside to moderately behind the anus
and just inside pneumostome. Mantle collar often without lobes or
lap>. but a glandular extension onto pallial roof present in some taxa.
Genital system variable, but apical sections typically as follows:
ovotestis of two closely appressed, variously subdivided, teardrop-
shaped lobes lying just above stomach apex; hermaphroditic duct a
simple, uncoiled tube entering laterally into carrefour swelling;
talon with globular head on a short stalk entering apex of carrefour;
albumen gland elongate-ovate, deeply indented by loops of intestine
and head of spermatheca. Prostate-uterus partly fused to fused, free
oviduct short to long, uterus with different glandular zones. Sper-
matheca with lower shaft expanded, upper shaft more slender after
tapering, head ovate, expanded, lying imbedded in base of albumen
gland above pallial cavity apex. Vas deferens a slender tube entering
penis complex near insertion of penial retractor muscle to well below
muscle insertion. Penis complex with a penis-derived epiphallic sec-
tion in many taxa, interior of penis with long and irregular pilasters,
often with a short vergic papilla. Epiphallic section, when present,
with slightly different textured pilasters. Entire complex slender and
tuhelike. Penial retractor muscle arising from diaphragm, inserting
on head of penis or epiphallic section, without complexities of inser-
tion. Interior of free oviduct and base of spermathecal shaft with
irregular folds in the few examined. Digestive and free muscle sys-
tems agreeing with endodontid and charopid patterns. No data are
recorded on the nervous system.
The family unit Punctidae is used here in a re-
stricted sense, not as the umbrella for the endodontoids
as suggested by Climo (1969a, 1971a, 1978). Pending
anatomical confirmation, I include the Punctidae or
Punctinae of Northern Hemisphere workers, Laominae
in the sense of Suter (1913) and Gabriel (1930), and
Paralaomidae of Iredale (1941a, 1944). As a rough es-
timate, about 159 named taxa from the southern parts
of Australia, New Zealand, subantarctic islands,
Holarctic, and scattered African localities would be in-
cluded. Only a very few taxa have had other than jaw
and radula data recorded in the literature, and it is
quite probable that some of the included species and
genera will prove to be charopids and some taxa as-
signed to the charopids will prove to be punctids. Par-
ticularly when shell sculpture becomes reduced, a
common pattern in the Austro-Zelandic taxa, assign-
ment to a family group on shell features alone becomes
extremely hazardous.
The anatomy o{Punctum Morse (1864) was illus-
trated and discussed by Baker (1927, pp. 227-228, pi.
16, figs. 8-12) then reprinted in Pilsbry (1948, pp.
641-643, fig. 349a-f). Data on the shell sculpture for-
mation was given by Solem (1977b, pp. 150-152, figs.
4-6, 11-12). Anatomical data are set forth here on
Punctum (fig. 24a-b), the New Zealand Laoma
leimonias (Gray, 1850) (fig. 25a-h), Western Austra-
lian Paralaoma aprica Iredale, 1939 (fig. 26a-c), and
Tasmanian rr Paralaorna" coesa (Legrand, 1871) (fig.
27a-d). These extralimital taxa are illustrated to show
the essential unity of structure within the Punctidae
and to provide data for comparing charopid patterns.
FIG. 24. Partial anatomy ofPunctum minutissimum (Lea). Cedar
bog, Woodburn Road, 4 miles southwest of Urbana, Champaign
County, Ohio. E. Keferl! X-20-1969. FMNH 151102: a, pallial region;
b, apical genitalia with albumen gland removed. No scale lines pre-
pared. (CW).
To my knowledge, no other taxa have been illustrated
in equivalent detail. An anatomical description of
Laoma leimonias follows to provide detailed compara-
tive data.
Laoma leimonias (Gray, 1850).
Description of soft part*. Foot and tail slender, rounded pos-
teriorly, not tapering noticeably. Sole undivided, without corruga-
tions i fig. 25d). Pedal grooves high on foot, suprapedal very much
smaller than pedal, uniting above tail. Pedal groove passing around
tail with a distinct constriction on top of tail. No protruding caudal
horn, but top of tail indented, with subpedal region extending pos-
teriorly. No middorsal groove. Slime network very inconspicuous.
Body roundly truncated anteriorly.
Body color in preservative yellow white, ommatophores black.
Surface of pallial roof and visceral hump with frequent black speck-
les, less frequent white marks.
Mantle collar (MC) elongated (fig. 25a), with bluntly rounded
anterior edge. Basal margin with long intrusion of mantle gland onto
pallial roof, corresponding with position of large left anterior mantle
lobe (MA I at parietal-palatal angle. Anus (A) opening at inner edge
of mantle collar (fig. 25b), slightly anterior to external urinary pore
(KX). Urinary chamber (LK) a sharply defined groove that is im-
mediately united with the V-shaped anal channel passing forward in
the mantle collar.
Pallial region (fig. 25a-b) very elongated, extending P/i-2
whorls apically from aperture. Basal Vi whorl with glandular exten-
sion of mantle collar (MG). longest where major palatal lamella is
formed and reabsorbed. Kidney (K) U-shaped, pericardia! and rectal
branches equal in length. Rectal arm of kidney lapping completely
under hindgut (HG) and extending onto parietal margin of pallial
cavity (fig. 25c). Thus, rectal arm lies on both parietal and upper
palatal margins of pallial cavity. Pericardia! arm of kidney appears
more slender in Figure 25a, but solely because pericardium lies un-
derneath kidney. Hindgut (HG) paralleling parietal-palatal margin
well past apex of pallial region. Loop of intestine (I) with apical base
KD ^-^\ Q
""" "p^~"
HV
2 mm
1mm
x
^
^r 2
GD
FIG. 25. Anatomy of Laoma Icimonian iGrayi. Herekino Gorge, Kaitaia, Northland, North Island, New Zealand. L. Price! XI-1962. FMNH
135401: a, pallia! region; b, detail of pneumostomal openings; c, detail showing extent to which kidney <K) overlaps hindgut (HG); d, extended
foot showing pedal grooves and lack of a caudal horn; e, genitalia; f, junction of hermaphroditic duct (GO) and talon <GT); g, detail of uterine
structure; h, internal structure of penis and epiphallus. Scale lines as marked. (PS).
55
56
SOLEM: ENDODONTOID LAND SNAILS
of kidney extended above it for a considerable distance. Ureter (KD)
sigmurethrous, arms without any lung roof visible between them,
not expanded on any portion. When first removed, apical rectal arm
of ureter hidden by parietal extension of kidney, but in Figure 25a
this is stretched outward. Heart (H) comparatively large, about half
as long as kidney, lying partly under pericardia! arm of kidney,
reaching pallial roof only on basal margin of whorl. Principal pulmo-
nary vein (HV) narrow, reaching mantle gland, without noticeable
secondary venation.
Ovotestis (G) composed of 1-2 clumps of finger-like lobes (fig.
25c), buried in apical section of digestive gland above stomach-
intestine reflexion, lighter in color than digestive gland. Hermaph-
roditic duct (GD) slender at first, gradually becoming a thick, lumpy,
muscular tube, slightly iridescent in tone. Duct narrows abruptly
just at base of albumen gland then reflexes up along albumen gland
(GG) before entering duct of talon (GT). Latter a slender tube with
expanded head (fig. 25f), entering head of prostate-uterus without
differentiated carrefour. Prostate (DG) composed of a few relatively
large acini fastened to wall of upper uterine chamber. Lower part of
prostate squeezed between uterine sections. Uterus (UT) three-
chambered: upper section (UT,) a very thin-walled tube, opening into
a second chamber (fig. 25g) with glandular walls and two or more
low, rounded pilasters (UT 2 ). Lower chamber (UT,) barely distin-
guished externally from second chamber, a thin-walled sac with
lamellar plates on the wall.
Vas deferens (VD) arising from narrowed base of prostate, a
slender tube becoming moderately expanded, then narrowed again
before inserting laterally on epiphallus (E) between arms of a narrow
U-shaped stimulator (fig. 25h). Vas not tightly bound to penioviducal
angle. Epiphallus weakly differentiated from penis, a short swollen
tube with a U-shaped stimulator and a narrow neck before joining
the penis proper (P). At apex of penis lies a high muscular stimulator
(PP), ovate in outline and attached at both ends, its apical tip lying
between the arms of the epiphallic stimulator (fig. 25h).
Below penial stimulator, there are two weak "pilasters" caused
by folding of the penis wall and very faint spiral pustulations. Bas-
ally, penis tapering to a very slender tube before entering short at-
rium ( Y). Gonopore a short slit behind and below right ommatophore.
Free oviduct (UV) long with thick muscular wall, a glandular
collar separating its base from spermathecal pore. Spermatheca (S)
long, expanded head lying partly in albumen gland and partly in
digestive gland, lying above pallial cavity. Lower part of sper-
mathecal stalk slightly larger than oviduct, thin-walled with distinct
longitudinal pilasters, narrowing at base of prostate-uterus to a very
slender tube, partly buried in acini of prostate. Vagina (V) very
short, not differentiated internally from atrium and base of penis.
Buccal mass of average length, high, strongly tapered an-
teriorly, generative sac small. Buccal retractor massive, not split,
inserting at about posterior third of buccal mass in U-shaped fan.
Esophagus initially large in diameter, then slender for length of
pallial cavity, inserting slightly behind midpoint of buccal mass.
Interior of esophagus with weak longitudinal pilasters, becoming
fainter apically. Esophagus extends about 1 A whorl past head of pal-
lial cavity, entering stomach along lower parietal margin of shell.
Stomach very large, reaching surface of parietal and upper palatal
margins, extending apically for slightly more than one whorl. Intes-
tine reflexing along columellar margin to lower margin of kidney
base, looping up along kidney base to just below hindgut, reflexing
posteriorly and diagonally downward to base of stomach, then loop-
ing up to parietal-palatal margin and running forward as hindgut.
Hindgut follows parietal-palatal margin along edge of pallial cavity
to anus. Loop of intestine abutting kidney projects into kidney, which
extends outside of intestine apically, thus separating body wall from
pallial roof.
Salivary glands white, lying alongside esophagus, touching me-
dially, near posterior end. Salivary ducts straight, entering laterally
on base of esophagus.
Digestive gland extending from apex of soft parts to apex of
pallial cavity, reduced in region of stomach, distinctly darker in color
than ovotestis.
Free muscle system simple: buccal retractor massive, not split
anteriorly, uniting with tail fan to form columellar retractor about
two whorls above aperture. Right ommatophoral retractor passing
between penioviducal angle. Rhinophoral retractors very short, unit-
ing with ommatophorals just before union with tail fan. Entire free
muscle system short and massive, extending apically less than 2M>
whorls from aperture.
(Based on 4 adult examples from south side of Herekino Gorge,
10 mi. southwest of Kaitaia, Northland, New Zealand (35 17' S, 173
12' E), FMNH 135401. Collected by L. Price, November 1962.)
Of the dissected species, Punctum minutissimum
(Lea, 1841) andParalaoma aprica (Iredale, 1939) are in
the 1.5-2.0 mm. size range; Laoma leimonias (Gray,
1850) is a very high-spired shell, about 2.3 mm. in
diameter and 2.7 mm. high with 7 whorls; and "Para-
laoma" coesa (Legrand, 1871) is the largest species,
about 3.5-4.0 mm. in diameter, with 4Vz whorls. As a
systematic comment, Punctum and Paralaoma may
well be synonyms, but "Paralaoma" coesa probably
does not belong to the same genus. There are
nomenclatural problems in dealing with this species.
The very long, free oviduct and penis of Laoma is re-
lated to the greatly increased whorl count of that
species and has no special systematic significance.
The basic anatomical features of the Punctidae
are: (1) the bicuspid lateromarginal teeth with minute
accessory cusps (fig. 12b-e), long basal plates, and for-
ward pointing of the rounded cusps; (2) the ovotestis of
two teardrop-shaped lobes that may be appressed (figs.
24b, 25e, 26c) or modestly separated (fig. 27b) and are
subdivided to various extents (compare figs. 24b and
27b with Baker, 1927, p. 16, fig. 8); (3) the bilobed
kidney of approximately equal lobe length, with (fig.
27a) or without (figs. 24a, 25a, 26a) lung roof visible
between the ureter arms; (4) the slender penis complex
with simple insertion of the penial retractor muscle
that internally has a penis-derived epiphallic section
followed by vergic papilla and smooth regular to ir-
regular pilasters; and (5) normally rather slender ter-
minal female genitalia with tapering lower sper-
mathecal stalk, internally with free oviduct, vagina,
and spermatheca sharing smooth pilaster sculpture.
The Charopidae contrast in having: (1) tri cuspid
laterals and marginals with rectangular basal plates, a
simplified interrow tooth support system during feed-
ing, and upward-slanting pointed cusps (figs. 13-14);
(2) the ovotestis consisting of one to two clumps of sev-
eral long follicles that lie parallel to the shell axis (figs.
90i, lOOb-c, g); (3) a generally bilobed kidney in which
the lobes may be equal in length (figs. 39c, 90a), the
rectal much longer (figs. 53a, e, 90e), or infrequently
the pericardial lobe is longer (fig. lla), and there may
(fig. 39c) or may not (fig. 90e) be lung roof visible be-
tween the arms of the ureter; (4) the penis complex
normally is short and fat (fig. 90b, f, i), with simple to
complex insertion of the penial retractor muscle, the
penis frequently with a verge, complex stimulators and
stimulator pockets, a vas deferens-derived epiphallus
in many taxa, and the lower penis area without
sculpture; and (5) normally thick terminal female
SYSTEMATIC REVIEW
57
FIG. 26. Partial anatomy of Paralaoma aprica Iredale. Station
WA-98, north end of townsite reserve, Nangeenan, west of Merredin,
Western Australia. A. Solem and L. Price! 11-22-1974. FMNH
182222: a, pallial region; b, postapical genitalia; c, apical genitalia.
Scale lines as marked. (EL).
genitalia with very thick spermathecal base that
rather abruptly narrows to the mid stalk, internally
often with very prominent pilasters that may be corru-
gated (fig. 9d) or smooth but very sharply defined (fig.
87c).
The Endodontidae contrast to both in having: (1)
bicuspid laterals and marginals with the marginal ec-
tocones split (Solem, 1976b, p. 91, fig. 54a-f), rectangu-
lar basal plates with interrow support system (Solem,
1976b, p. 89, fig. 52d), and rather sharp elevation of the
pointed cusps (Solem, 1976b, p. 89, fig. 52b); (2) the
ovotestis of multiple clumps angled to the shell axis
(Solem, 1976b, p. 372, fig. 163c); (3) the kidney with at
most a weak rectal lobe developed and only a primary
ureter that opens at the front edge of the rectal kidney
lobe near the posterior of the pallial cavity (Solem,
1976b, p. 459, fig. 195a); (4) a slender penis that nor-
mally has only two simple longitudinal pilasters, never
a verge or vergic papilla, only rarely a penis chamber-
derived epiphallic section (Solem, 1976b, p. 396, fig.
171b-c, h); and (5) very thin terminal female genitalia
without special internal sculpture and with highly var-
iable insertion of the spermathecal stalk (Solem,
1976b, p. 84, fig. 48a-c). In addition, the complete sep-
aration of the prostate and uterus and apparent ab-
sence of a sperm packet in the Endodontidae are major
differences from either of the other families.
The changes outlined above are noncorrelated, and
their different states in the three families indicate that
the families represent three separate patterns of diver-
sification. These differences are greater than those
seen between limacacean families and thus merit rec-
ognition as family units. Whether the punctid radula is
correlated with specialized feeding habits as opposed to
those of the Charopidae and Endodontidae is unknown.
The Punctidae are only marginally present in the
Pacific Island area, with a scattering of records in
Hawaii, a single record from the mountains of Tahiti,
and a new species, Punctum polynesicum, described
here from the Austral Islands. Thus, no attempt is
made at revising the generic classification or discuss-
ing detailed patterns of distribution. Reference of these
Pacific specimens to Punctum is based strictly on con-
chological similarity in the absence of preserved
specimens. The distinction between Paralaoma and
Punctum may not be maintained in the future, but the
shells of the Pacific Island taxa are far more similar to
the Holarctic Punctum than to the Austro-Zelandic
Paralaoma. In view of the limited material, no diag-
nosis is presented of the genus Punctum.
Genus Punctum Morse, 1864
Type species. Helix minutissima Lea, 1841, by
original designation.
There are about 50 sets of Punctum from various
Hawaiian Islands in the Bernice P. Bishop Museum
which may be lumped temporarily as Punctum horneri
Ancey (1904, p. 66, pi. 5, figs. 11-12), but no material
has been reported previously from Polynesia. A single
specimen (BPBM 145298) from Station 865, Mt. Aorai
Trail, Tahiti, collected between 5,600 and 6,300 ft. ele-
vation, can be identified as a Punctum, but is too frag-
mentary for any attempt at giving it a specific name.
The high spire and narrow umbilicus of the fragment
combine with the finer and less protracted sculpture to
place it in Punctum rather than Striatura.
Euconulus (Monoconulus) conoidea H. B. Baker
(1941, pp. 214-215), the only non-Hawaiian Pacific Is-
land Euconulus known to date, was taken at the same
collecting station. Presence of these otherwise Ha-
waiian and Holarctic taxa in Tahiti at high elevations
could be explained by the deus ex machina of accidental
transport on the feathers of migratory birds.
More abundant material of quite different species
from the Austral Islands is described below as
Punctum polynesicum. Although somewhat atypical in
whorl count and heavy sculpture, it is nearer to
Punctum than any other genus known to me. Only
dead specimens were available. Measurements are
summarized in Table XII.
58
SOLEM: ENDODONTOID LAND SNAILS
UT
FIG. 27. Anatomy of "Paralaoma" coesa (Legrand). Station TA-38, Circular Head, near Stanley, North Tasmania. L. Price! XI-8-1966.
FMNH 154974: a, pallial region; b, genitalia; c, interior of penis; d, interior of free oviduct and spermatheca. Scale lines as marked. (NB).
Punctum polynesicum, new species. Figure 28a-c.
Diagnosis. Shell minute, diameter 0.95-1.02 mm. (mean 0.99
mm.), with 3V8-3%+ rather tightly coiled whorls. Apex and spire
moderately and evenly elevated, body whorl descending more
rapidly, spire protrusion about V4 body whorl width (0.30-0.34 mm.),
H/D ratio 0.563-0.597 (mean 0.580). Apical whorls IMs, sculpture of
about 15 fine spiral cords, whose interstices are 3-4 times their
width. Postnuclear whorls with narrow, rounded, strongly protrac-
tively sinuated, prominent radial ribs, 72-83 (mean 77.3) on the
body whorl, whose interstices are about twice their width until end of
body whorl where ribbing becomes crowded. Ribs/mm. 23.2-25.9
(mean 24.9). Microsculpture of fine radial riblets, 2-3 between each
pair of major ribs, crossed by equally fine spiral riblets. Umbilicus
shallow, broadly V-shaped, regularly decoiling, contained 3.33-3.44
times (mean 3.37) in the diameter, margins rounded. Sutures deep,
SYSTEMATIC REVIEW
59
TABLE XII. - RANGE OF VARIATION IN PUNCTUM, DISCOCHAROPA, AND MICROCHAROPA.
NUMBER OF
SPECIMENS
NAME
Punctum
polynesicum 6
Discocharopa
aperta(Mlldf.) 310
Microcharopa
mlmula
RIBS
77.3(72-83)
110.6(77-148)
RIBS/MM.
24.9(23.2-25.9)
26.9(18.0-40.4)
126.6(117-136) 36.5(32.3-40.5)
HEIGHT DIAMETER H/D RATIO
0.58(0.56-0.61) 0.99(0.95-1.02) 0.580(0.563-0.597)
0.59(0.48-0.74) 1.36(1.18-1.84) 0.438(0.341-0.615)
0.48(0.43-0.53) 1.07(0.95-1.13) 0.449(0.400-0.469)
WHORLS UMBILICUS D/U RATIO
polyn. 3 1/4(3 1/8-3 3/8) 0.29(0.28-0.30) 3.37(3.33-3.44)
aperta 3 l/2-(3 1/8-4 1/4) 0.54(0.43-0.92) 2.58(2.00-3.21)
APICAL
CORDS
MAJOR
RADIALS
SPIRE
ELEVATION
BODY WHORL
WIDTH
SP/BWW
0.08(0.07-0.10) 0.32(0.30-0.34) 0.255(0.216-0.316)
0.07(0.03-0.15) 0.40(0.35-0.44) 0.170(0.086-0.286)
mimula 3 l/4-(3 1/8-3 3/8) 0.40(0.35-0.46) 2.68(2.43-3.05) 17.8(16-20) 0.05(0.04-0.07) 0.32(0.28-0.34) 0.186(0.118-0.289)
whorls strongly rounded, with slight lateral compression above
periphery and on basal margin. Aperture circular, inclined about 30
from shell axis. Color light yellow-horn, without darker markings.
Punctum polynesicum is the smallest species yet
described within the genus. Its very prominent radial
ribbing and rather wide umbilicus separate it from
most species. Punctum vitreum H. B. Baker, 1930, is
perhaps nearest in strength of ribbing, but is a much
larger shell with a lower H/D ratio.
Description. Shell minute, with slightly more than 3V4 rather
tightly coiled whorls. Apex and spire moderately and evenly ele-
vated, body whorl descending much more rapidly, H/D ratio 0.589.
Apical whorls I'/i, sculpture partly eroded. Postnuclear sculpture
and microsculpture as in diagnosis, 83 ribs on the body whorl. Um-
bilicus broadly V-shaped, regularly decoiling, contained 3.35 times
in the diameter. Sutures and whorl contour typical, aperture shape
and inclination as in diagnosis. Height of holotype 0.59 mm., diame-
ter 1.02 mm.
Holotype. Austral Islands: Raivavae, Station
622, Ahuoivi Point. Collected dead under hau leaves by
Y. Kondo, D. Anderson, and C. M. Cooke, Jr., on Au-
gust 9, 1934. BPBM 147099.
Range. Tubuai and Raivavae, Austral Islands.
Paratypes. Raivavae: west side Vaiaunana Pe-
ninsula (Station 587), 100 yd. inland at 200 ft. eleva-
tion, under dead leaves (three specimens, BPBM
146725). Tubuai: southwest of Murivai (Station 698),
300-400 yd. inland under Barringtonia trees (two
specimens, BPBM 147679).
Remarks. Without preserved material for dissec-
tion, classification of this species will remain uncer-
tain. The presence of endodontoid reticulated micro-
sculpture between the major ribs, rather than the
transverse growth wrinkles seen in Striatura (Solem,
1977b, p. 151, fig. 1) has dictated my placing this
species in Punctum. In several respects P. polynesicum
agrees more with Striatura. The latter averages fewer
whorls (3 l /4-3%) and has a wider umbilicus than does
Punctum with its 3%-4% whorls and generally nar-
rower umbilicus. Apical sculpture in Striatura is gen-
erally more prominent and narrower, whereas in
Punctum it is wider and lower. The low whorl count
and fine apical sculpture of P. polynesicum thus appear
somewhat anomalous.
Neither genus has been reported from South
Pacific Islands, although both live in Hawaii. A typi-
cally sculptured Punctum is reported from Tahiti
above, but this species is very different in form, color,
shell thickness, and ribbing from P. polynesicum. Mi-
crocharopa could be confused on the basis of size, but
that species has radically different apical sculpture
(see fig. 4, p. 13), a much lower H/D ratio (mean 0.449),
and many more, much finer radial ribs (117-136 on the
body whorl).
An alternative possibility is that this species rep-
resents the outermost Polynesian representative of the
Charopidae and is related to Sinployea. I consider it
much more probable that it is a Punctum accidentally
introduced by migratory birds. In the absence of dissec-
tions, no subgeneric reference would be meaningful.
FAMILY CHAROPIDAE
Small to large endodontoids, apical sculpture of shell very di-
verse, postapical whorls primitively with major radial ribs and a
microsculpture of radial riblets and secondary spiral elements, most
sculpture features composed of periostracal elements only. Many
taxa with sculpture secondarily reduced to lost. Whorl counts and
shell shape highly variable. Umbilicus widely open to closed. Color
monochrome to brightly tessellated. Apertural barriers evolved in-
dependently in several lineages, most species without trace of bar-
riers. Visceral hump and shell greatly reduced to almost lost in a few
taxa. Foot of animal undivided, prominent pedal and suprapedal
grooves uniting above tail, in some taxa forming a caudal foss or
mucus pore. Radula with tricuspid central slightly to distinctly
smaller than normally tricuspid laterals, rarely bicuspid or with only
one cusp. Marginals frequently tricuspid, sometimes with ectoconal
splitting. Basal plates of central and marginal teeth rectangular,
interrow support system well developed, latter absent on marginals.
Cusps of major teeth sharp and tending to point upward with ec-
tocones elevated much more than in Endodontidae. Jaw of separated,
squarish plates in smaller species, becoming partly to completely
fused in larger taxa. Pallial complex usually with a distinctly bilobed
kidney, relative size of lobes quite variable among species, plus a
closed secondary ureter that normally opens just inside pneumo-
stome. Some taxa with only a partial secondary ureter that opens far
60
SOLEM: ENDODONTOID LAND SNAILS
h H
FIG. 28. a-c, Punctum polynesicum, new species. Station 622,
Ahuoivi Point, Raivavae, Austral Islands. Holotype. BPBM 147099.
Scale line equals 1 mm. (LH).
posteriorly in the pallial cavity, and/or rectal kidney lobe greatly
reduced. Mantle collar normally without prominent lobes or laps, a
glandular extension onto pallial roof found in some taxa. Genital
system highly variable. Ovotestis typically of one or two clumps of
long follicles that lie parallel to the outer wall of the body cavity and
occur sequentially. Some taxa with the ovotestis broken up into a
series of lobes that are perpendicular to the outer wall of the body
cavity and widely separated from each other, a few taxa with follicles
shortened and resembling those of the Punctidae. Hermaphroditic
duct a simple unkinked, usually iridescent tube entering usually
laterally onto apex of carrefour. Talon a finger-like projection or
swollen head on a short stalk. Albumen gland irregular in shape,
deeply indented by loops of intestine and head of spermatheca. Pros-
tate and uterus fused with partial common lumen, uterus with sepa-
rate and sequential glandular sections. Free oviduct short to long,
thin, base of spermatheca and upper vagina normally swollen. Sper-
mathecal shaft tapering abruptly, head above apex of pallial cavity,
an expanded oval, to elongate oval structure. Vas deferens slender,
normally entering a clearly differentiated epiphallus, often through
a complex valvular pore. Many taxa with epiphallus secondarily
compressed into penis or absent. Penis usually fat, at least in upper
section, not usually long and slender, lower part often a thin tube.
Interior of penis frequently with a well-developed verge, circular
ridges, stimulatory pilasters, and pocket stimulators. Penis retractor
arising from diaphragm, usually very short, inserting in simple to
complex fashion on the penis or epiphallus. Interior of lower female
tract simple to with very large longitudinal simple to corrugated
pilasters, openings to free oviduct and/or spermathecal shaft varying
from simple pores to with complex valves. Penis, so far as observed,
enervated from right cerebral ganglion. Digestive and free muscle
system typically endodontoid.
The Charopidae, as rather broadly defined here,
includes the Phenacohelicidae, Otoconchinae, Flam-
mulinidae, Rotadiscinae, Amphidoxinae, Dipnelicidae,
Hedleyoconchidae, and Pseudocharopidae of authors.
In terms of species, it is the dominant land snail group
of New Zealand (Powell, 1976; Climo, 1969a-b, 1970,
1971a-b, 1978), New Caledonia (Franc, 1957; Solem,
1961), parts of Australia (Tasmania, Victoria, less
abundant in New South Wales, South Australia,
southwestern Western Australia, sparse in the North-
ern Territory, well represented in the rain forest areas
of Queensland; Iredale, 1937a-c, 1939, 1941a-b;
Burch, 1976; Solem, in preparation C), Lord Howe Is-
land (Iredale, 1944), Norfolk Island (Iredale, 1945), and
Kermadec Islands (Iredale, 1913). The Charopidae is
only sparsely represented in Indonesia (Solem, 1958),
New Guinea (Solem, 1958, 1959b, 1970a), Solomon Is-
lands (Solem, 1960), and New Hebrides (Solem, 1959a).
Charopids occur with varying abundance on the is-
lands surveyed in this report.
Extralimital to the Pacific Basin, species of
charopids are common to dominant in southern Africa
(Connolly, 1939; Solem, 1970c), were common on St.
Helena (Solem, 1977c), and Notodiscus (Solem, 1968c)
is found on the Subantarctic Islands (Kerguelen, Pos-
session, Heard, Marion, Amsterdam). There is a mod-
est Juan Fernandez radiation (Ohdner, 1922) and
many scattered descriptions and records from southern
South America, the Andes, and Central America. Two
species occur in western North America as far north as
Idaho and Montana (Pilsbry, 1948).
Including the taxa discussed and described below,
approximately 646 named species are included in the
SYSTEMATIC REVIEW
61
Charopidae. This is a very conservative estimate of ac-
tual extant diversity because Climo (personal com-
munication) has large numbers of undescribed New
Zealand taxa, a rain forest patch survey of Queensland
and Northern New South Wales revealed approxi-
mately 30 undescribed taxa (Burch, 1976), and my own
1974 and 1976-1977 field collections in southwestern
Western Australia produced at least 15 new species
(Solem, in preparation C). In contrast, several years of
intensive fieldwork in Victoria by Brian Smith and
associates from the National Museum of Victoria have
confirmed the described diversity and greatly extended
ranges but have not produced new taxa. It is thus im-
possible to present an accurate estimate of diversity in
the Charopidae at the present time, except to state that
the currently recorded taxa are significantly less in
number than the extant taxa.
A breakdown on estimated endodontoid species
numbers is given in Table XI. The grand total of 1,494
described and collected but undescribed taxa contrasts
with the species estimate given by Boss (1971, p. 108)
of only 502 species of endodontoid land snails. A
critique of Boss's assumptions and procedures was in-
cluded in Solem (1978b).
Because this monograph deals with taxa from only
one area of species abundance, it is inappropriate to try
for a worldwide revision of higher categories. More im-
portant, data in the literature is not adequate to make
higher level taxonomic decisions, and only part of the
material needed to work out these problems is avail-
able, dissected, and illustrated. Illustrations and de-
scriptions needed to justify most synonymization of
family names and to emphasize the broad distributions
involved are published, but work on the Western Aus-
tralian taxa (Solem, in preparation C), including the
Dipnelicidae, is omitted at this time.
Where possible, actual type species have been dis-
sected and illustrated, but in taxa such as Paralaoma
(figs. 26-27) this was not possible. Anatomical illus-
trations and descriptions ofCharopa coma (Gray, 1843)
(figs. 9-lO),Phenacohelixpilula (Reeve, 1852) (fig. 11),
Flammulina zebra (Le Guillou, 1842) (fig. 29), Am-
phidoxa marmorella (Pfeiffer, 1845) (fig. 30),
Stephanoda binneyana (Pfeiffer, 1847) (fig. 31), and
Pseudocharopa lidgbirdi (Etheridge, 1889) (fig. 32),
based on new dissections, are presented here. Litera-
ture data on the anatomy of Rotadiscus (Baker, 1927,
pp. 228-230, pi. 16, figs. 13-20) and the Otoconchinae
(Climo, 1969a, fig. 2C; 197 la) allow handling of all the
included subfamily names except the Hedleyoconch-
inae. This Australian group, fairly common in New
South Wales and Queensland, with a possible relative
on Lord Howe Island (Iredale, 1944, pp. 317-318), has
not been dissected, but probably is another example of
shell sculpture reduction and spire elevation in an ar-
boreal taxon. External body features, jaw, and radula
were figured by Hedley (1888, fig.; 1889, pi. 15) and
copied by Pilsbry (1893-1895, pi. 9, figs. 27-29). These
features are typically charopid in aspect.
Australian and New Zealand workers have vari-
ously used the names Flammulinidae, Charopidae,
Phenacohelicidae, and Otoconchidae for subgroups of
this complex (Gabriel, 1930; Iredale, 1937a et seq.;
Climo, 1969a-b, 1970, 1971a-b, 1978; Powell, 1976).
In large part this was based on early work of Henry
Suter, which was summarized in his magnificent
monograph (Suter, 1913). The ancient question as to
the significance of the caudal pore or foss was discussed
above and well summarized by Climo (1969a). It has no
major taxonomic utility. Climo (1969a) focused on the
presence or absence of an epiphallus as a significant
subfamily key character. The externally easily recog-
nizable epiphallus ofCharopa (fig. 9b, d) does contrast
with the situation in Phenacohelix (fig. lie, e) where
the insertion of the penial retractor has shifted to the
epiphallus-vas deferens junction, rather than inserting
on the penis-epiphallus junction as in Charopa. In
Flammulina (fig. 29b, e-j) the situation is complicated
by visceral hump reduction and drastic increase in
whorl profile cross-sectional area. In Flammulina
zebra, the genotype, the epiphallus has been compacted
back into the penis (fig. 29e-h) and is thus completely
undetectable from external view (fig. 29b). In other
taxa referred to Flammulina (Climo, 1969a, fig. 2A)
and the Otoconchinae (Climo, 1969a, fig. 2B-C; 1971a,
fig. 1E-F) greater compaction has taken place in the
region of the prostate and uterus, whereas the penis
and epiphallus region are maintained as separate en-
tities.
Anatomical descriptions of the family units' type
genera follow to provide comparative data with the
later dissections and to record structural variations for
consideration by future revisors of extralimital taxa.
The descriptions follow the same sequence as the illus-
trations Charopa coma (Gray) (figs. 9-10), Phe-
nacohelix pilula (Reeve) (fig. 11), Flammulina zebra
(Le Guillou) (fig. 29),Amphidoxa marmorella (Pfeiffer)
(fig. 30), Stephanoda binneyana (Pfeiffer) (fig. 31), and
illustrations only of Pseudocharopa lidgbirdi
(Etheridge) (fig. 32). Discussion of anatomical varia-
tion in that species is deferred to a review of the pat-
terns in Lord Howe Island taxa (Solem, in prepara-
tion). Nomenclatural details and a shell description of
Amphidoxa marmorella are included because so little
has been recorded about this species.
Charopa coma (Gray, 1843).
Description of soft parts. Foot and tail slender, rounded pos-
teriorly, but not tapering. Sole undivided and without any corruga-
tions (fig. 9a). Pedal grooves not particularly high on foot, suprapedal
less distinct than pedal, both grooves uniting over tail. No caudal
horn or foss. Middorsal groove weak. Slime network rectangular,
fairly inconspicuous. Head extended well in front of foot, which is
squarely truncated anteriorly. Ommatophores long.
Body color in preservative light yellow-white, ommatophores
black.
Mantle collar (MC) rather narrow (fig. 9e f) with a large glandu-
lar extension (MG) encroaching on lung surface. Pneumostome (LP)
masked by quite small left anterior mantle lobe (MA) at parietal-
palatal angle. Anus (A) opening slightly anterior to external ureteric
62
SOLEM: ENDODONTOID LAND SNAILS
pore (KX) and sharing single opening with pneumostome (LP). Uri-
nary chamber (LK) a narrow groove angling past anus.
Pallial region (fig. 9e) varying in length from %-l whorl, elon-
gated, anterior portion filled with mantle gland (MG). Lung roof
clear, except for occasional white speckles near kidney tip. Kidney
(K) U-shaped, rectal arm equal in length to pericardia!, but dis-
tinctly more slender, lapping under hindgut. Hindgut (HG) parallel-
ing parietal-palatal margin past top of pallial cavity, slender. Kidney
abutting on loop of intestine (I). Ureter (KD) sigmurethrous, not
expanded at any portion, both arms compreseed between lobes of
kidney, then bordering hindgut to external ureteric opening (KX).
Heart (H) proportionately large, much shorter than kidney. Prin-
cipal pulmonary vein following edge of kidney, then running forward
to edge of mantle gland. No conspicuous secondary venation.
Ovotestis (G) embedded in apical lobe of liver (fig. 9b), variable
in form and spacing, consisting of a few finger-like lobes, often split
one or more times. Individual lobes with slender pink rods. Mass of
ovotestis may be concentrated at one point or divided into two masses
separated by part of digestive gland. Hermaphroditic duct (GD)
straight, thick-walled, iridescent pink, sometimes with one or more
knobs on surface, narrowing slightly to radically just before reaching
albumen gland (GG) and entering midway on talon (fig. 9b, GT).
Latter a blind pouch buried in albumen gland, head enlarged and
pink, duct slender, passing straight into head of prostate-uterus.
Prostate (DG) of large rectangular glandular lobes pressed against
surface of uterine sector. Uterus (UT) an extremely thin-walled cir-
cular tube partly filled with spongy tissue opening through a con-
stricted collar into a thick-walled, white, expanded glandular pore
with smooth inner walls except for a single large pilaster. Muscle
fibers pass from uterine base to columellar retractor.
Vas deferens (VD) arising from narrowing of prostate along
glandular section of uterus, first part expanded, then continuing as
narrow tube along free oviduct and vagina to penioviducal angle,
then up penis to epiphallus (E). Epiphallus short, thick-walled, ex-
panded, internally (fig. 10) with two longitudinal pilasters, refiexed
and entering penis (P) through a small verge (fig. 9d, PV). Penial
retractor muscle (PR) arising on diaphragm, attached to apex of
penis in a U-shaped fan. Penis expanded, internally with thick pus-
tulose glandular lining extending from apex to near base which ter-
minates in a thickened collar. Verge small, with terminal pore, not
developed in juveniles. Below level of verge, a blind pocket with
smoothly muscled walls develops. This "penial caecum" (PC) is
strictly internal and not visible on outside of penis. Glandular collar
considerably thicker than rest of penis interior and gives false ap-
pearance of being a verge when atrium (Y) opened.
Length of free oviduct (UV) highly variable, apparently corre-
lated with length of pallial region. In figured individual it is ex-
tremely short; in others almost equal to vagina in length. Base of free
oviduct constricted by a glandular papilla with central pore (UVO).
Spermatheca (S) with very slender duct, bound to surface of prostate,
passing into visceral mass above pallial cavity and expanding into a
distinct head. Albumen gland also in mass of digestive gland and
intestinal loop above pallial cavity. Vagina (V) long, swollen me-
dially, with a prominent tubercle, or set of tubercles (VS), located
between oviducal pore and the several longitudinal pilasters that
extend to atrium (Y). Latter short, thin-walled, opening into gono-
pore, a transverse slit located just behind and above right
rhinophore.
Free muscle system simple. Tentacular retractors (TP) uniting
with tail fan (FR) well in front of union with buccal retractor (BR).
Point of union between ommatophoral (TER) and rhinophoral (TVR)
retractors variable, usually about midway along distance to tail fan.
Right ommatophoral retractor passing between penioviducal angle.
Buccal mass (B) short and high (fig. 9g), with very slender
generative sac. Buccal retractors (BR) insert as U-shaped fan about
Vt of way from posterior end. Esophagus (BE) very long and slender,
originating high on buccal mass only slightly behind midpoint and
well in front of buccal retractor insertion. Interior of esophagus with
longitudinal ribs. Stomach originating just above pallial cavity, ex-
tending apically for % of a whorl and with reflexion of stomach at
point well below first lobe of ovotestis. First arm of intestine follow-
ing lower inner margin of whorl to base of kidney, looping below
cephalic aorta and across base of kidney, then reflexing for less than
1 k length of stomach, rising to upper parietal-palatal margin and
passing forward as hindgut.
Digestive gland extending from apex of soft parts to just above
intestinal loop, lower parts finger-like because of the huge stomach.
Main artery, granular white in color, empties primarily into stomach
and secondarily into recurved lower loop of intestine.
Salivary glands (OG) elongate, situated on sides and top of
esophagus, closely associated with prostate-uterus in body cavity.
Salivary ducts (OD) straight, inserting on each side at base of
esophagus.
(Based on FMNH 135420, Waiwera-Puhoi Road, North of Auck-
land, North Island, New Zealand, A. Solem! February 10, 1962. Sev-
eral adults and juveniles.)
Phenacohelix pilula (Reeve, 1852).
Description of soft parts. Foot long and slender, tapering
gradually to a sharply rounded posterior margin. Sole undivided,
without corrugations. Pedal grooves very high on foot, suprapedal
distinctly less impressed than pedal, both grooves uniting over tail.
Caudal horn a very long, tapering projection (fig. lib, CH). Middor-
sal groove absent. Slime network very weak, barely visible. Head
extending only slightly in front of foot, which is bluntly rounded
anteriorly.
Body color in preservative yellow-white, with faint grayish tinge
to upper part of tail and caudal horn, ommatophores reddish black.
Dark color markings on pallial roof visible through shell.
Mantle collar (MC) narrow (fig. lla), with moderate glandular
intrusion (MG) onto pallial roof. Pneumostome without clearly de-
fined lappets. Anus (A) opening above and slightly anterior to exter-
nal ureteric pore (KX), urinary chamber (LK) a shallow groove lead-
ing past anus and fading out toward mantle collar margin.
Pallial region (fig. lla) rather short and compact. Lung roof with
numerous scattered black speckles and much less frequent white
patches. Kidney (K) bilobed, pericardia! arm much longer than rec-
tal, 3.5 times length of pericardium (H). Base of kidney abutting on
loop of intestine. Rectal arm of kidney lapping under hindgut, less
than half length of pericardial arm. Ureter (KD) sigmurethrous,
pericardia! arm slightly expanded over that along hindgut (HG). A
clear section of lung roof visible between arms of ureter. Neither
ureter nor hindgut expanded during anterior passage toward
pneumostome. Hindgut (HG) paralleling parietal-palatal angle well
past apex of pallial cavity. Heart (H) short, angled to plane of
hindgut. Principal pulmonary vein (HV) simple, extending forward
to edge of mantle gland intrusion, without conspicuous secondary
venation.
Ovotestis (fig. lie, G) buried in apical lobes of digestive gland
above stomach-intestine reflexion, consisting of one or two clumps of
pinkish iridescent finger-like alveoli joined by slender ducts in a "Y"
leading to main hermaphroditic duct (GD). Latter a slender duct
tapering gradually toward an enormously swollen portion, about 1 A
of duct length, shortly before approach to albumen gland, then be-
coming an extremely slender tube before joining talon (fig. lid, GT),
which is buried in albumen gland (GG). Latter a blunt, finger-like
projection lying buried in visceral hump above pallial cavity and
composed of quite large acini with a yellow-orange tinge. Basal por-
tion of hermaphroditic duct reaching base of albumen gland, refiexed
apically along surface before entering laterally upon enlarged head
of talon, which is partly buried within albumen gland. Narrowed
shaft of talon running to head of prostate-uterus, no expanded car-
refour visible. Prostate (GD) of much larger, lighter colored acini
than those of albumen gland, fastening to thin-walled, upper uterine
chamber. This is a flat, smooth tube gradually tapering from a nar-
row apex to a broad base where the second uterine chamber (UT 2 ), a
glandular walled structure, expands. Lower part of uterus a thin-
walled chamber with internal lamellar plates, opening into a bul-
bously expanded free oviduct.
Vas deferens (VD) arising as groove from base of prostate, partly
hidden by third uterine chamber, passing down as a thin-walled tube
SYSTEMATIC REVIEW
63
to penioviducal angle (to which it is lightly bound by fibers), reflex-
ing along penis and becoming progressively larger in diameter and
more muscular before joining epiphallus laterally. Penial retractor
(PR) inserting directly on head of epiphallus (E), a short, solid,
globose tissue mass arising on diaphragm. Epiphallus an expanded
muscular tube entering penis (P) through a smooth muscular verge
(fig. lie, PV). Walls of penis with weak crenulated pilasters and
pustulations basally, apical portion with glandular collar just below
tip of verge. Externally, position of glandular collar and verge indi-
cated by swelling in penis-epiphallus. Atrium (Y) shortened and
leading to gonopore located below and a little behind right om-
matophore.
Free oviduct (UV) a short, globosely expanded structure narrow-
ing slightly to point of union with spermathecal stalk. Spermatheca
(S) with very slender stalk, partly buried between acini of prostate,
passing apically to small expanded head lying above apex of pallial
cavity in visceral mass together with albumen gland and talon. Va-
gina (V) much longer than free oviduct with a very thin-walled ac-
cessory pocket that was badly torn in all dissected specimens. The
exact relationship of the vagina, oviduct, and accessory vaginal
pocket was not determined.
Buccal mass rather short, strongly curved on top, generative sac
very small. Buccal retractors insert in U-shaped fan about '/-.< of way
from posterior of buccal mass. Esophagus inserting nearly at mid-
point of upper surface of buccal mass, very slender and thin-walled,
extending through visceral hump past apex of pallial cavity. At this
point, the esophagus expands into a funnel shape, entering stomach
yet allowing room for positioning of intestinal loops. Esophagus en-
tering stomach on lower inner whorl margin just above apex of pal-
lial cavity. Stomach occupies nearly entire whorl, grossly expanded
to cover outer and upper margins, extending apically to point below
"Y" of hermaphroditic duct, intestine reflexing along base of whorl,
about midway between inner and outer margin. Intestine running
apically to apex of pallial cavity, looping across outer whorl margin
and abutting on kidney, reflexing backward and downward along
expanding margin of stomach, then looping upward on outer wall to
parietal-palatal margin and passing forward as hindgut
Digestive gland extending from tip of soft parts to apex of pallial
cavity, greatly reduced in region occupied by stomach.
Salivary glands slender, white, lying along and surrounding
esophagus. Salivary ducts entering buccal mass-esophagus junction
laterally.
Free muscle system simple. Columellar retractor very large,
buccal retractor uniting with tentaculars shortly before joining main
tail fan.
(Based on FMNH 135421, Church Road, Kaitaia, Northland,
New Zealand, L. Price! October 1962, several adults.)
Flammulina zebra (Le Guillou, 1842).
Description of soft parts. Foot and tail about equal in length to
shell diameter, truncated anteriorly, rapidly tapering from about 1
mm. behind visceral stalk posterior. Sole undivided longitudinally,
with weak transverse corrugations extending from sides of foot.
Pedal grooves high on foot, suprapedal slightly weaker than pedal,
united above tail, no caudal horn, or middorsal groove present. Slime
network irregularly rectangular, relatively weak. Head and om-
matophores without unusual features. Gonopore on right side of
head, directly behind ommatophore.
Body color dark gray on head and tail, lighter between pedal
grooves, soles and sides of foot yellow- white.
Mantle collar (fig. 29a, MC) without lobes or laps, no glandular
extension onto pallial roof. Anus (A) opening slightly anterior to
external ureteric pore (KX). Urinary chamber (LK) not dif-
ferentiated.
Pallial region (fig. 29a) extending about V:> whorl apically, quite
broad, lung roof without color markings. Kidney (K) bilobed, short,
quite wide. Ureter (KD) thick, arms sharply angled, with angle ap-
proaching 80. Heart (H) half length of kidney. Principal pulmonary
vein (HV) weakly branched, stopping well short of mantle collar.
Hindgut (HG) slender, reaching parietal-palatal margin opposite
point where ureter reflexes, then running forward to anus.
Ovotestis two large clumps of palmately clavate alveoli buried in
digestive gland above stomach apex. Clumps with central orienta-
tion perpendicular to plane of coiling, laterally fanning from center.
Hermaphroditic duct narrow, variously kinked and coiled, extending
to side of prostate, reflexed upward and straightened to angled union
with carrefour (fig. 29c-d). Albumen gland large, of slender acini,
tightly packed around carrefour region. Talon (GT) very short, with
circular head, buried in albumen gland. Carrefour (X) with only
slight enlargement from talon shaft (fig. 29c-d). Prostate (DG) of
comparatively few acini along sides of uterus, exact structure of pas-
sage from carrefour to vas deferens not determined. Possibly a nar-
row part of the uterine lumen serves to transfer sperm. Uterus (UT)
grossly enlarged, twisted, with thick glandular walls. Vas deferens
(VD) thicker for first V:i of length, then narrowed for last %, entering
apex of epiphallus next to point where penial retractor muscle in-
serts, tightly bound to penis surface (fig. 29b). Epiphallus compacted
into upper penial chamber (fig. 29e-i), with complex valvular en-
trance from vas deferens (fig. 29i-j). Penial retractor (PR) very short,
scarcely a separate muscle zone. Penis (P) almost circular, about 1.3
mm. long, apex (fig. 29e) with compacted epiphallus opening through
a corrugated verge (fig. 29e-f, PV), lower portion with a complex set
of pilasters, highly modified from standard charopid pattern. Atrium
(Y) long and slender, thin-walled, without internal sculpturing (fig.
29b).
Free oviduct (UV) short, twisted, fusing indistinctly with lower
organs. Spermatheca (S) expanded basally, shaft following side of
uterus, curving up and around carrefour region, head partly next to
exposed section of prostate, partly next to albumen gland, oval in
shape. Vaginal and lower penis area fused into a common chamber.
Free muscle system short, characterized by lack of fusion be-
tween muscles. Right ommatophoral retractor (TER) not passing
through penioviducal angle, arising laterally from columellar mus-
cle. Right rhinophoral retractor (TVR) arising from columellar mus-
cle just inside origin of ommatophoral retractor.
Buccal mass shortly oval, proportionately very large, with small,
generative sac. Buccal retractors split, each half inserting laterally
near posterior margin of buccal mass, inserting at apex of columellar
muscle. Esophagus short, not enlarged, arising from middle of buccal
mass upper surface. Stomach starting expansion before apex of pal-
lial cavity, occupying '/z whorl, then reflexing to intestine, which
loops down and forward along lower whorl margin, looping inward
and upward along side of pericardium and pericardial kidney, then
making a curved U-loop backward and inward before angling for-
ward as hindgut to parietal-palatal margin. Hindgut slanting down
from parietal-palatal margin at point of ureter reflexion.
Digestive glands typical. Salivary glands lying on top and on
either side of esophagus, partly fused medially.
(Based on FMNH 153217, Okuti Valley, Banks Peninsula, near
Christchurch, New Zealand, F. Climo! March 7, 1965. Three dis-
sected adults.)
Amphidoxa marmorella (Pfeiffer, 1845).
Helix marmorella Pfeiffer, 1845, Proc. Zool. Soc. London, 1845, p.
125- Island of Juan Fernandez; Pfeiffer, 1847, Monog. helic. viv.,
1, p. 66; Albers, 1850, Die Heliceen, Isted., p. 110; Pfeiffer, 1852,
Syst. Conchol. Cab., I, 19(2), pp. 129-130, pi. 88, figs. 17-19
(plate issued in 1850); Reeve, 1852, Conchol. Icon., Helix, pi. 113,
fig. 649.
Helix (Amphidoxa) marmorella Pfeiffer, Tryon, 1887, Man.
Conchol., (2), 3, p. 46, pi. 20, fig. 54.
Amphidoxa (Amphidoxa) marmorella (Pfeiffer), Pilsbry, 1893,
Man. Conchol., (2), 9, pp. 39-40, pi. 7, figs. 10-12.
Amphidoxa marmorella (Pfeiffer), Odhner, 1922, Nat. Hist., Juan
Fernandez and Easter Island, 3, pp. 229, 230-231, figs. 9-10
(radula and jaw).
Description of shell. Shell small, with slightly more than 3
rapidly enlarging whorls (fig. 30a). Apex slightly emergent, penul-
timate and body whorl flatly coiled, H/D ratio 0.550. Apical whorls
FIG. 29. Anatomy of Flammulina zebra (Le Guillou). Okuti Valley, Banks Peninsula, near Christchurch, New Zealand. F. Climo! HI-7-1965.
FMNH 153217: a, pallial region; b, postapical genitalia; c-d, views of the hermaphroditic duct-talon-carrefour junction; e, interior of penis
chamber; f-h, detail of vas deferens-epiphallus-penis junction; i-j, detail of vas deferens-epiphallus junction. Scale lines equal 1 mm. foro-6, e;
other figures greatly enlarged. (CW).
64
HG
KO
PR
1 mm
FIG. 30. Anatomy of Amphidoxa marmorella (PfeifTer). 500 m. elevation, northeast slope, Portazuelo, Masatierra, Juan Fernandez. B.
Malkin! IV-7-1962. FMNH 135424: a, apical view of shell prior to dissection; b, basal view of shell; c, pallial region; d, genitalia; e, interior of
penis. Scale lines as marked. (PS).
65
66
SOLEM: ENDODONTOID LAND SNAILS
1%, sculpture of 20 protractively angled large radial ribs over a mi-
croreticulated lattice visible only under 96 x magnification. Remain-
ing whorls with protractively sinuated, lamellar radial ribs, 32 on
the body whorl, whose interstices are more than 8 times their width.
Ribs elevated greatly by periostracal extensions. Microsculpture a
lattice of microradial and microspiral riblets, the former slightly
stronger. Sutures deep, whorls rather flatly rounded above evenly
rounded periphery and base. Color light yellow-white with very
vague reddish flammulations, waved, becoming strongly retractive
on base of shell, apparently not extending into umbilicus. Umbilicus
narrow, V-shaped, regularly decoiling, contained 8.3 times in the
diameter. Aperture large, subcircular, slightly flattened laterally
above periphery, inclined about 30 from shell axis. No apertural
barriers. Height of shell 1.64 mm., diameter 2.99 mm.
Description of soft parts. Body completely retracted within
shell (fig. 30b), soft parts relatively hardened and difficult to dissect.
Foot relatively short and broad, rounded behind, apparently not ta-
pering, truncated anteriorly. Sole undivided longitudinally, with
rather strong transverse corrugations near tail, corrugations coming
from slime network. Pedal grooves deep, high on foot, suprapedal as
marked as pedal, both grooves uniting over tail. Because of contrac-
tion, presence or absence of middorsal groove could not be deter-
mined. Slime network a sharply defined irregularly rectangular
network. Ommatophores short and chunky.
Body color yellow-white, mantle collar brownish. Eyespots small
and black, upper part of stalk with black speckles.
Mantle collar (fig. 30c, MC) thick, glandular, without any man-
tle gland extension onto lung roof. Pneumostome masked by bulges
in mantle collar, but no distinctive lobes present. Anus (A) opening
at inner edge of mantle collar, a slight groove running from it to
outer edge of collar.
Pallial region (fig. 30c) extending about '/n whorl, shape and
positioning of structures probably altered by extreme contraction of
animal. Lung roof clear, without dark speckling except in region of
ureter and pericardium and a very few along hindgut near anus.
Kidney (K) bilobed, main lobe pericardial, with a narrow tongue
extending along hindgut (HG). Main lobe twisted in front of heart
(H) extending on both sides of ureter (KD) tip. Base of kidney extend-
ing outside of intestinal loops along outer whorl edge (partly an
artifact of contraction?). Entire first loop of intestine and part of loop
to hindgut lying next to kidney. Ureter (KD) following anterior mar-
gin of pericardial kidney arm upward and slightly backward into
small pocket between pericardial arm of kidney and hindgut then
ending in a pore pointing apically. No groove leads toward pneumo-
stome nor is there any trace of a secondary ureter. Heart (H) rather
large, constricted by retraction. Principal pulmonary vein (HV) fol-
lowing edge of kidney and tip of ureter, then passing forward to
mantle collar with no conspicuous lateral venation.
Ovotestis (fig. 30d, G) two clusters of finger-like lobes, some
bifurcated, imbedded in digestive gland above stomach, a Y-fork col-
lecting tubule attaching them to hermaphroditic duct (GD). Latter
extending along stomach at basal margin of whorl, a thick, nodular,
opaque, iridescent tube becoming slender near albumen gland and
running to stalk of talon. Albumen gland (GG) finely textured, oc-
cupying visceral area between stomach base and apex of pallial cav-
ity inside of intestinal loops. Head of spermatheca (S) lying in angle
between prostate head and albumen gland base. Talon (GT) long
with slender, slightly tapering head and broader base that is buried
in albumen gland. Hermaphroditic duct entering talon below apex of
slender stalk. Talon opening into apices of prostate and uterus. Pros-
tate (DG) of large acini opening into a closed duct that is partly
enfolded by uterus (UT1. Lower part of prostate squeezed by ex-
panded uterus and opening into slender vas deferens (VD). Uterus
(UT) a slender, thin-walled tube above, opening into a grossly ex-
panded chamber with thicker glandular walls and weak traces of
ridging.
Vas deferens (VD) a very slender tube passing to penioviducal
angle, around right ommatophoral retractor, then up to head of penis
(P). Penial retractor (PR) arising on diaphragm, inserting on head of
epiphallus (E). Vas deferens inserting laterally on epiphallus, which
contains longitudinal pilasters surrounding the vas pore and leading
into the verge (fig. 30e, PV). Penis proper bulbous above, tapering to
a slender duct before joining vagina (V) to form atrium (Y). Inter-
nally (fig. 30e) penis with a large apical verge (PV) whose tip is
bifurcate; a large circular muscular collar (PP,) separated into two
parts; a broad, thin pilaster (PP 2 ) tapering to the slender basal stalk;
and an apparently doughnut-shaped pilaster (PP :1 ) that is partly
folded over upon itself in the available specimen. Atrium (Y) short,
rather large in diameter.
Free oviduct (fig. 30d, UV) short, only slightly narrowed from
uterus, internally with glandular walls and vague pilasters. Sper-
matheca (S) with basal part of shaft slightly expanded, tapering
along surface of uterus and prostate, expanded head lying in angle
between prostate and albumen gland. Vagina (V) very large, ex-
panded, internally with broad longitudinal glandular pilasters that
continue into atrium (Y). No special pilasters or collars mark sper-
mathecal or oviducal terminations. A small vaginal diverticulum
(VC) is a blind pocket opening between two major pilasters that
continue up into spermatheca.
Buccal mass ovoid, strongly contracted, generative sac rather
large. Buccal retractor inserting in U-shpaed fan about Vy of way
from posterior end, not split. Esophagus entering top of buccal mass
slightly behind midpoint, a rather thick tube with longitudinal pilas-
ters following inner margin of whorl past apex of pallial cavity and
entering stomach very shortly past that point. Stomach extending
about '/4 whorl, expanded to fill parietal and upper palatal walls
almost immediately, narrowing about Vh of way from apex, the nar-
rowing resulting in both a descension from the parietal-palatal angle
and basal narrowing. At apex, stomach reflexes downward and turns
near basal margin before joining intestine. At stomach-intestine
junction, ducts of digestive gland enter on each side. Intestine follow-
ing basal-palatal margin of whorl forward to base of kidney (fig. 30c,
K), looping across kidney base, nearly touching hindgut, looping di-
rectly downward, then backward and upward as hindgut to par-
ietal-palatal margin, finally following that margin to anus. Extreme
sharpness of intestinal looping may be result of contraction.
Digestive gland extending from apex of soft parts to apex of
pallial cavity; a broad band along lower palatal wall between intes-
tine and outer part of stomach, reduced in amount in area of intesti-
nal loops and albumen gland. Openings into stomach-intestine junc-
tion very prominent.
Salivary glands white, finely textured, lying lateral to esopha-
gus, touching above. Ducts of salivary glands entering buccal mass
on each side of esophageal insertion.
Free muscle system short and massive. Buccal retractor nearly
as broad as long when contracted. Right ommatophoral retractor
passing between penioviducal angle, uniting with rhinophoral re-
tractor just before joining tail fan. Tentacular retractors joining tail
fan subapically and laterally, buccal retractor joining medially and
apically.
Jaw fragmented during mounting, composed of about 20 delicate
plates that are weakly striated and do not strongly overlap.
Radula with 75 rows. Central tooth very small, with weak side
cusps. Lateral teeth large, tricuspid, endoconal size gradually in-
creasing during shift to marginals, ectocones on outer marginals
split.
(Based on FMNH 135424, northeast slope at 500 m. elevation,
above Portazuelo, Masatierra, Juan Fernandez, B. Malkin! April 7,
1962. One adult specimen.)
Stephanoda binneyana (Pfeiffer, 1847).
Description of soft parts. Foot narrow and rather elongated, not
tapering posteriorly, bluntly rounded behind (fig. 31f). Pedal grooves
deeply and equally impressed, high on sides of foot. Sole undivided
longitudinally, but strongly corrugated perpendicular to body axis,
such corrugations being continuations of lateral grooves below pedal
grooves. Tail with middorsal groove, bell-shaped in cross section,
sloping down to point where pedal grooves unite. No caudal foss or
horn developed. Slime network fine, irregular, oblong to rectangular
patches.
Body color in preservative pale yellow-white, sides of head and
mantle with gray markings, ommatophores black.
Mantle collar (MC) thick with tapering edge (fig. 31a). Pneumo-
SYSTEMATIC REVIEW
67
FIG. 31. Anatomy of Stephanoda binneyana (Pfeiffer). Rio Cisnes (44 45' S, 72 W), Prov. Aysen, Chile. L. E. Pena! 11-1961. FMNH 135428:
a, pallial region; b, genitalia; c, free muscle system; d, interior of terminal genitalia; e, detail of epiphallus-penis junction; f, rear portion of tail;
g, detail of anterior mantle cavity; h, detail of hermaphroditic duct-talon-carrefour junction; i, lobe of ovotestis. Scale lines as marked, detail
sketches greatly enlarged. (PS).
stome (LP) masked by small left anterior mantle lobe (MA). Edge of
collar just anterior of pneumostome has a weak thickening forming
an accessory mantle lobe (AMA). Anus, excretory pore, and
pneumostome sharing common opening (fig. 31g). Anus (A) slightly
anterior to external ureteric pore (KX). Urinary chamber (LK) a
shallow groove running past anus and continuing (although less
sharply defined) nearly to edge of mantle collar.
Pallial region (fig. 31a) elongated, but in view of contracted state
no accurate estimate of length possible. Lung roof with variable col-
oring, speckled with widely scattered black and white flakes to
nearly coal black. Kidney (K) short, elongately triangular with a
narrow, hooked basal extension abutting hindgut (HG). Base of kid-
ney against loop of intestine. Ureter (KD) slender, sigmurethrous,
flaring slightly midway along hindgut, arms of ureter widely sepa-
rated by lung roof. Heart (H) slightly more than '/2 length of kidney.
Principal pulmonary vein (HV) angling toward pneumostome,
nearly reaching mantle collar, without conspicuous side branches.
Ovotestis (G) imbedded in digestive gland above stomach-
intestine reflexion, consisting of 8 sets (4 shown in fig. 31b) of
clumped palmately clavate alveoli (fig. 31i) strung linearly along
very thin hermaphroditic duct (GD). Ovotestis distinctly lighter in
tone than digestive gland. Collecting tubule of hermaphroditic duct
68
SOLEM: ENDODONTOID LAND SNAILS
very thin, bilobed. Main section a round muscular tubule, slightly
iridescent in tone, narrowing just before base of albumen gland (GG),
reflexing abruptly and ascending to apex of bilobed talon (GT) (see
fig. 31h). Albumen gland elongately ovate, blunt-tipped, lying com-
pletely above pallia! cavity, very fine-textured. Talon (GT) buried in
albumen gland, quite small, white. Duct from talon opening into
head of prostate-uterus without differentiated carrefour. Prostate (D)
equal in width to upper portion of uterus, individual acini much
larger than those of albumen gland. Lower part of prostate enveloped
in basal sections of uterus. Uterus (UT) tripartite. Upper section
narrow, sacculated, equal in width to prostate. Basal '/n of uterus
surrounded by a hollow, smooth-walled, white-colored chamber with
very fine texture. Basal section of uterus expanded into a chamber
with lamellar walls.
Vas deferens (VD) originating from groove of prostate enfolded
by second uterine chamber, a narrow tube passing down to peni-
oviducal angle, then almost immediately into epiphallus (E). Latter
a much larger muscular tube, first half swollen with strong pilasters
on one wall, then narrowing to about 4 times diameter of vas def-
erens. Epiphallus entering penis (P) just below attachment of penial
retractor (PR), which is short, quite thick, and originates on dia-
phragm. Penis rather short, apical half bulbous, tapering toward
atrium. Internally (fig. 31d-e) with rather heavy and complex pilas-
ters. A single large lamellar plate (PP) starts just below epiphallic
pore, curves up and around it, then passes longitudinally down wall
opposite insertion of penial retractor. A second longitudinal pilaster
(PP,) arises below origin of major pilaster and tapers gradually to-
ward atrium. A series of circular minor pilasters complete the ar-
mament.
Free oviduct (UV) short and muscular, bulbous, internally (fig.
3 Id) with rugose pilasters. Spermatheca (S) of long type, expanded
head in visceral mass above pallial cavity and cephalic aorta, stalk
very slender. Vagina (V) long, with twisted, bilobed accessory vagi-
nal appendix (VC). Main lobe (fig. 31b, d) with muscular walls, a few
longitudinal pilasters, and an intricately reticulated stimulatory pad
(VS). Secondary lobe (VC) a bulbous, tapering, thin-walled organ,
the expanded head with many lamellar plates. Stalk of secondary
lobe and duct of vagina with weak longitudinal pilasters. Main ac-
cessory lobe of vaginal appendix bound to right side of tail fan muscle
(fig. 31c, VRM) slightly less than halfway from union with right
tentacular retractors and insertion on foot.
Free muscle system (fig. 31c) simple. Buccal retractor (BR) at-
taching on posterior Va of buccal mass, in U-shaped fashion, but not
split until very tip, and joining tail fan (FR) at site of columellar
attachment (CR). Ommatophoral retractor (TER) and rhinophoral
retractors (TVR) unite a short distance back and almost immediately
fuse with tail fan. Right ommatophoral retractor passing between
penioviducal angle. Tail fan split slightly posterior to junction with
tentacular retractors (TR). Penis base bound to body wall, accessory
lobe of vagina bound to right part of tail fan (VRM).
Buccal mass rather short and muscular, generative sac promi-
nent. Esophagus entering top of buccal mass between 1 A and '/a way
from posterior end, large and muscular, with heavy longitudinal
ridges anteriorly, tapering and becoming thin-walled to apical end of
pallial cavity. Stomach a very large thin-walled sac extending less
than 1 whorl apically, along parietal-palatal margin and occupying
much of space in visceral hump. Reflexing as intestine along lower
inner margin of whorl, running back to apex of pallial cavity, curv-
ing up and abutting on kidney base, looping back diagonally down-
ward for '/2 length of stomach, then reflexing forward diagonally as
hindgut. Hindgut not reaching parietal-palatal margin until just
above start of pallial cavity, following margin to anus and only
slightly expanded in lower part of pallial cavity. Salivary glands
paired, lying on each side of esophagus, touching above and below.
Ducts very slender, inserting on each side of posterior esophageal
base. Extreme duct coiling and nearness of glands to buccal mass
probably result of extreme contraction.
Digestive gland extending from top of pallial cavity to apex of
soft parts (which do not reach shell apex). Lower section with intesti-
nal loops finger-like. Minor collecting tubules pass through spaces
between branches of ovotestis clumps. Major duct empties into base
of stomach.
(Based on FMNH 135428. Rio Cisnes, Aysen, Chile, L. E. Pena!
February 1961. Four dissected adults.)
All of these taxa agree in basic structures of the
genitalia, pallial region, radula so far as observed, and
muscle system. Their differences relate to visceral
hump and compaction changes in the case of Flam-
mulina and the Otoconchinae, multiple radiations be-
tween terrestrial and arboreal habitats (shell sculpture
reduction, mucus pore intensification), and species rec-
ognition factors in the terminal genitalia. The latter
changes are many and bewildering, since 15 to 30 en-
dodontoids from a single locality is normal in New Zea-
land. The variations in penis interior structure illus-
trated by Climo (1969b, figs. 25A, 28D, 29B, C, 30E;
1970, figs. 16B, 20D, 2 1C) can be directly compared
with the type of sympatric variations reported below.
Continuation of the Otoconchinae as a separate
subfamily, based on the multiple changes in structure
resulting from visceral hump reduction, can be jus-
tified and is accepted here. It is less strongly
differentiated than the other subfamilies recognized in
this report, and rigorous structural analysis may result
in determining that the Otoconchinae is a grade in
structure rather than a clade.
The remaining New Zealand Charopidae are not
amenable to subfamily distribution at this stage.
Champa, Phenacohelix, and Flammulina agree in all
essential details of structure. They are reasonably
closely related. The many illustrations of Climo
(1969a-b; 1970) do suggest possible subgroupings, but
until details of vergic, epiphallic, and lower female
tract structures are described and illustrated, no real
decisions are possible. Although the New Zealand
charopids have radiated extensively and confusingly,
their basic structures seem rather coherent. Retention
within an admittedly rather broad subfamily Charo-
pinae is suggested pending detailed revisions.
Because most of the family names used for the
Australian charopids have been based on New Zealand
genera, the great variation found in these taxa will not
be discussed further. Study of Western Australian taxa
with primitive features is in progress (Solem, in prep-
aration C). The family name Dipnelicidae Iredale
(1937b, pp. 22-23) will be considered in that report.
The structural variety in the Australian taxa seems
greater than that reported for the New Zealand
species, but this is only a preliminary impression.
The family Pseudocharopidae Iredale (1944, p.
312) was proposed by the phrases "An extraordinary
development of beautiful small shells is indicated by
this family name. ... I was thinking of allotting them
to the Flammulinidae, which they slightly resemble.
... to save further confusion the above name (Pseudo-
charopidae) is utilized." Dissection of Pseudocharopa
lidgbirdi (Etheridge, 1889) (fig. 32a-c) shows a typical
pallial cavity configuration (fig. 32a), with bilobed
kidney, complete secondary ureter, and essentially un-
SYSTEMATIC REVIEW
69
branched pulmonary vein. The genitalia (fig. 32c) is
unusual only in having a single kink in the hermaph-
roditic duct (GD), small prostate, shortened terminal
female organs, the epiphallus shifted in relation to the
penial retractor muscle, and the interior of the penis
(fig. 32b) with high, smooth pilasters in the lower
penis, rugose low pilasters in the upper penis. Because
the shell of this species has a reduced visceral hump,
most of the unusual features can be interpreted as sec-
ondary. The basic features are those of the Charopinae.
KD
FIG. 32. Anatomy of Pseudocharopa lidgbirdi (Etheridge). Sta-
tion 20, northeast slope of Mt. Lidgbird, 1,500 ft. elevation, south end
Lord Howe Island. L. Price! IX-1963. FMNH 127977: a, pallial re-
gion; b, interior of penis complex; c, genitalia. Scale lines as marked;
b greatly enlarged. (NB).
Amphidoxinae Thiele (1931, p. 578) is based on
one of the Juan Fernandez Islands species, A mphidoxa
marmorella (Pfeiffer, 1845). A single, retracted, nearly
adult specimen (fig. 30a-b) was dissected. The
genitalia (fig. 30d-e) has the typical charopid aspect,
with the addition of a vaginal caecum (VC), shortened
epiphallus (E), and internally shows the verge (PV),
ring pilasters (PPi), and pocket stimulator (PC) seen in
Champa (fig. 9d) and Sinployea (figs, 43e, 57c). The
only unusual features are the absence of a secondary
ureter (fig. 30c) and the retention of a separate pros-
tate. Despite this, the basic structures of Amphidoxa
and Charopa are amazingly similar. At least the
structures of Amphidoxa show how the Charopinae
could be derived from more generalized stocks.
Traditionally, the South American genus
Stephanoda has been associated with Amphidoxa. Dis-
section of Stephanoda binneyana (Pfeiffer, 1847)
showed profound differences. The pallial region (fig.
31a) has the short rectal lobe found in Phenacohelix
(fig. lla), a complete secondary ureter (KD) that opens
(KX) (fig. 31g) just posterior of the anus (A). In the
genitalia (fig. 31b, d-e, h-i) the ovotestis consists of
several follicle bundles (fig. 31b, i), the talon (GT) is
reduced (fig. 31h), the prostate-uterus is typical, but
the slender base of the spermatheca (S) (fig. 31b) and
fat free oviduct (UV) again agree with Phenacohelix
(fig. lie). The tail of the animal (fig. 31f) has the typi-
cally conservative form found in Charopa (fig. 9a) and
all Pacific Island taxa. The most unusual features are
in the terminal genitalia, with a prominent vaginal
caecum (VC) (fig. 31b), long vagina (V), and short penis
(P). Internally (fig. 31d-e) the penis has circular pilas-
ters and two large stimulators, but the epiphallus (E)
enters through a pore rather than a verge (fig. 31e).
There are irregular and complex pilasters within the
terminal female genitalia (fig. 31d). In the absence of
the verge and altered pilaster pattern, Stephanoda
shows considerable similarity to structures seen in
some of the non- verge bearing charopids illustrated by
Climo (1969b, 1970). These structures emphasize that
the charopids are a true Southern Hemisphere group
and that relationships are somewhat complex.
Comments on a few other extralimital taxa are
included under the appropriate subfamily discussions.
In this monograph I recognize five subfamilies in
the Charopidae: Otoconchinae Cockerell, 1893; Rota-
discinae Baker, 1927; Charopinae Hutton, 1884; Truk-
charopinae, new subfamily; and Semperdoninae, new
subfamily. Of these, the Otoconchinae is least sharply
differentiated, and the Charopinae is deliberately the
most broadly defined, pending adequate anatomical
revisions of the New Caledonian, New Zealand, Aus-
tralian, South American, and South African taxa. Un-
doubtedly the Charopinae will be subdivided further in
the future, but whether on a tribal basis or into several
subfamilies will depend upon the results of future in-
vestigations. A brief diagnosis of each subfamily fol-
lows.
70
SOLEM: ENDODONTOID LAND SNAILS
Subfamily Otoconchinae Tail and body greatly elongated, vis-
ceral hump reduced with shell a cap or fragment, head shortened in
relation to body length. Jaw fused, arcuate. Radula typical of
Charopinae. Pallial region modified Charopinae because of shorten-
ing, genital system compacted by head and visceral hump reduction,
basically Charopinae in structures. Otoconcha Hutton, 1884, and
Maoriconcha Dell, 1952, are the included taxa.
Subfamily Rotadiscinae Small to minute species with apical
sculpture of short spirally arranged segments (fig. 4a-e), visible op-
tically as spiral cords, postapical sculpture typically charopid. Jaw of
separate plates. Radula usually typical except for ectoconal marginal
splitting. Pallial region without to with a complete secondary ureter,
kidney weakly to evenly bilobed. Ovotestis of one to two clumps of
short to long follicles, talon short to long, not with circular head.
Terminal female system simple, sometimes with accessory caecum,
relatively thin. Epiphallus usually below penial retractor insertion,
short, opening into penis through a short conical verge or simple
pore, walls of penis with longitudinal pilasters. Rotadiscus Pilsbry,
1926, Radiodiscus Pilsbry & Ferriss, 1906, Radioconus H. B. Baker,
1927, Radiodomus H. B. Baker, 1930, Microcharopa, new genus, and
probably some undescribed genera from Western Australia are the
included taxa.
Subfamily Charopinae Small to large species with apical shell
sculpture of spiral cords, radial ribs, or secondarily smooth, postapi-
cal sculpture typically charopid, frequently secondarily reduced.
Apertural barriers developed in several lineages. Jaw of separate
plates in smaller taxa, fused partly to completely in large. Radula
generally with tricuspid laterals and marginals, but modified to
bicuspid or unicuspid laterals and sometimes with ectoconal splitting
on the marginals. Genitalia highly variable, epiphallus normally
present, terminal genitalia normally thick at least in part, penis
with or without verge, normally with complex stimulatory pilasters.
As a temporary measure, all charopid genera not specifically as-
signed to other subfamilies are lumped in the Charopinae.
Subfamily Trukcharopinae Small to medium-sized species
with apical sculpture of spiral cords, postapical sculpture strong to
secondarily reduced. Aperture with or without barriers. Jaw and
radula typically charopid. Genitalia unusual in lacking an epiphal-
lus. Vas deferens passing through the penial retractor muscle before
entering the penis either through a verge or pore. Interior of penis
with stimulatory pilasters of varied form. Trukcharopa,
Kubaryiellus, Russatus, Roimontis, Palikirus, Jokajdon, andPalline,
new genera from Micronesia, are the included taxa.
Subfamily Semperdoninae Medium-sized to large species with
spiral apical cords, typical to greatly reduced postapical sculpture.
Aperture with or without barriers. Jaw and radula without unusual
features. Pallial region typical, with some variation in kidney lobe
lengths. Genitalia unusual in the altered penis complex. Penial re-
tractor inserting on head of epiphallus, which is a coiled double tube,
outer wall thin muscle sheath, inner a thick glandular tube rolled
inward on one side, opening into penis through a simple pore. Penis
with an outer heavy muscle sheath of circular fibers that extends up
around the lower epiphallus, with an inner tube containing fine lon-
gitudinal pilasters above that usually coalesce into 3 glandular
pilasters below. Semperdon, Ladronellum, and Himeroconcha, new
genera from Micronesia, are the included taxa.
The affinities of the subfamilies are still uncer-
tain. The New Zealand Otoconchinae is an obvious
sluglike secondary derivative from the New Zealand
Charopinae. In structure it shows only minor dif-
ferences. The Trukcharopinae from Micronesia are de-
rivable from the typical Charopa-Sinployea stock by
loss of the epiphallus and specialization of the penial
surfaces. At present there are no Australian or New
Zealand structural equivalents known, but I would not
be surprised if relatives turn up in this area. The Sem-
perdoninae from Micronesia have a very different
penis structure than anything I have seen elsewhere.
Other features of their anatomy are compatible with
derivation from the Charopinae, and finding relatives
in a remote area of the family range would not surprise
me in the least.
In contrast, the Rotadiscinae have a relict (Idaho,
Arizona-Costa Rica, Trinidad, probably South
America), Fiji, Western Australia (Solem, unpublished
data) distribution. Their anatomy has a punctid aspect
in part to the ovotestis, the pallial region is more
primitive than that of the Charopinae, and its slender
terminal genitalia with some accessory organs is very
different from the Punctidae and most of the Charo-
pinae. This group may prove to be a relatively base
stock of the charopid radiation.
Further phylogenetic comments are postponed
pending completion of work on the Western Australian
radiation and additional work on New Caledonian and
Tasmanian taxa. Comments on relationships within
subfamilies are given below.
Subfamily ROTADISCINAE
A more expanded definition and discussion is post-
poned until completion of the Australian work. Inclu-
sion of Microcharopa in the Rotadiscinae is based on its
unique apical sculpture (fig. 4c-e) that is shared with
Radiodiscus (Solem, 1977b, pp. 150-154, figs. 7-8),
Rotadiscus, and undescribed Western Australian
material. Microcharopa could not be dissected. In shell
features it differs in its very small size and reduced
whorl count, finer and more crowded radial ribs, and
wide umbilicus. All of the New World genera are con-
siderably larger, more coarsely ribbed and generally
more variously umbilicated.
Genus Microcharopa, new genus
Shell minute, with less than 3M> tightly coiled whorls. Apex and
spire slightly elevated, body whorl descension a little more rapid.
Apical sculpture (fig. 4a-e) of short, twisted, open-ended ridges ar-
ranged in 16-20 spiral rows on top of a weakly and broadly undulat-
ing radially ridged surface. Postnuclear whorls with narrow, very
crowded, protractively sinuated radial ribs. Microsculpture of very
fine radials, finer spirals, and some weak secondary spiral cords.
Umbilicus widely open, cup-shaped, regularly decoiling, margins
weakly shouldered. Whorls almost evenly rounded, slightly com-
pressed laterally above periphery and on basal margin. Aperture
without barriers or heavy callus. Anatomy unknown.
Type species. Microcharopa mimula, new species.
Despite attempts on two different occasions to col-
lect material on Lami Ridge and near Sigatoka, Viti
Levu, no specimens could be found. Because only dead
material was collected previously, the anatomy of this
very unusual species could not be studied. Even under
100 x magnification with the light microscope, the
apex looks like it has only slightly wavy spiral cords,
more frequent in number than with most species, but
not very unusual in appearance. Scanning electron
microscope photographs at 300 x- 10,000 x magnifica-
tion (fig. 4a-e) show that the apical sculpture is very
complex. Instead of continuous spirals, there are rela-
tively short segments that are equal in width to the
SYSTEMATIC REVIEW
71
vague radial undulations (fig. 4a). Each individual
segment starts as a cord, twists up and over, then ends
as a cuplike structure (fig. 4d-e). These are in spiral
rows and are present except for the first quarter apical
whorl (fig. 4c), which seems to have low spiral cords
instead. Postnuclear sculpture (fig. 4b) shows no really
significant difference from that found in many
Charopidae.
Separating Microcharopa from other Pacific gen-
era is simple. The smallest Sinployea that has been
dissected, S. inermis lakembana, is more than twice the
diameter of Microcharopa, 2.36 mm. compared with
1.07 mm., and the smallest species assigned to
Sinployea, S. ellicensis nukulaelaeana, is 2.08 mm. in
mean diameter, only slightly less than twice the size.
Species of Vatusila are in the 1.65-1.88 mm. size
range, but these have very prominent apertural bar-
riers and much simpler apical sculpture. The only
genus in which the shells are approximately equal in
size is Discocharopa (p. 59). Populations of that taxon
average about 1.37 mm. in mean diameter and have
3V2-4 whorls, an even wider umbilicus, and, most im-
portant, very different apical sculpture (fig. 5a-c, p.
16).
The peculiar apical sculpture is shared with
Radiodiscus (see Solem, 1977b, p. 152, figs. 7-8), which
differs in having 3 3 /4-4Mi whorls, reaches 2.0-2.25 mm.
in diameter, and has about 21 ribs/mm, on the body
whorl, compared with 36.5 ribs/mm, in Microcharopa.
The narrower umbilici and larger sizes of Rotadiscus
and Radiodomus are easy recognition features of these
genera.
Only the one species, Microcharopa mimula, which
is widely distributed in the Lau Archipelago and on
Viti Levu, is known. The name Microcharopa refers to
the very small size of this species and its resemblance
in shape to the historical usage of the name Charopa.
Microcharopa mimula, new species. Figures 4a-e;
33a-c.
Diagnosis. Shell minute, diameter 0.95-1.13 mm. (mean 1.07
mm.), with 3Vs-3% rather tightly coiled whorls. Apex and spire
slightly to moderately and evenly elevated, body whorl descending
slightly more rapidly, spire protrusion about Ve-Vs body whorl width,
H/D ratio 0.400-0.469 (mean 0.449). Apical sculpture of 16-20
(mean 17.8) prominent, crowded spiral cordlike structures. Postnu-
clear sculpture of narrow, crowded, fine, sharply defined, protrac-
tively sinuated radial ribs, 117-136 (mean 126.6) on the body whorl,
whose interstices are 2-3 times their width. Ribs/mm. 32.3-40.5
(mean 36.5). Microsculpture of barely visible radial riblets, 1-2 be-
tween each pair of major ribs, equally fine spiral riblets, with occa-
sional traces of weak secondary spiral cording. Umbilicus broadly
open, cup-shaped, regularly decoiling, contained 2.43-3.05 times
(mean 2.68) in the diameter, margins weakly shouldered. Whorl con-
tours almost evenly rounded, slightly compressed laterally above
periphery and on basal margin, aperture circular, inclined about 10
from shell axis.
The similarities in size and shape to Discocharopa
are marked, but the spiral apical cording and absence
of a deeply recessed parietal lamella immediately
separate Microcharopa mimula from that genus. Other
barrier-free Fijian Charopinae are at least twice the
FIG. 33. a c, Microcharopa mimula, new species. Lami Ridge,
west of Suva, Viti Levu, Fiji. Holotype. BPBM 178490. Scale line
equals 0.5 mm. (MM).
size of M. mimula at 3 l /& whorls and show numerous
alterations in sculpture and shape.
Description. Shell minute, with 3 1 A rather tightly coiled whorls
that very slightly increase in width. Apex and spire slightly and
evenly elevated, body whorl descending moderately, H/D ratio 0.467.
Embryonic whorls slightly less than \ l k, sculpture of 18 crowded
72
SOLEM: ENDODONTOID LAND SNAILS
spiral riblets, sinuated where crossing low, broadly rounded, rela-
tively vague radial swellings. Postnuclear whorls with protractively
sinuated, crowded, narrow, lamellar radial ribs, 136 on the body
whorl, whose interstices are 1-2 times their width. Microsculpture
as in diagnosis. Sutures deep, whorls shouldered above, gently and
evenly rounded on outer margins. Umbilicus broadly open, saucer-
shaped, regularly decoiling, contained 2.70 times in the diameter,
margin weakly shouldered. Color very light yellow-brown. Aperture
circular, strongly rounded near periphery and at baso-umbilical
margin, inclined about 10 from the shell axis. Height of holotype
0.48 mm., diameter 1.02 mm.
Holotype. Fiji: Viti Levu, Station 47, Lami ridge,
5 miles west of Suva, at 5-150 ft. elevation. Collected
by Yoshio Kondo and C. M. Cooke, Jr., on July 24,
1938. BPBM 178490.
Range. Munia, Mothe, Wangava, Nayau, Na-
muka, and Yangasa Levu, Lau Group, and Viti Levu,
Fiji.
Paratypes. Viti Levu: Near Sigatoka, '/2 mile in-
land, under dead leaves, 50 ft. elevation (6 specimens,
BPBM 88524, BPBM 88562); Sigatoka, near base of
limestone cliff, Vz-lVfe miles from shore (1 specimen,
BPBM 87930); Nayavu, upper Wainibuka River, 14
miles inland, 190 ft. elevation (1 specimen, BPBM
87959); Lami Ridge, 5 miles west of Suva (Station 47),
limestone ridge, 5-150 ft. elevation (3 specimens,
BPBM 178490).
Lau Group: Munia, Vi mile inland (Station 65),
hillside in copra plantation at 600-900 ft. elevation (2
specimens, BPBM 179323); Mothe, central wooded
peak, on leaf mold at 590 ft. elevation (4 specimens,
BPBM 78585); Wangava, probably Station 27 (4
specimens, ex BPBM 166994); Nayau, Nauko (Station
43), hillside % mile inland at 250 ft. elevation (4 speci-
mens, BPBM 167231); Namuka, Matandolo (Station 2),
north-central point of island on hillside 500 ft. inland
at 65-75 ft. elevation (2 specimens, ex BPBM 166614);
Yangasa Levu, south end of island (Station 30), hill-
side, 150 ft. inland at 40 ft. elevation (2 specimens, ex
BPBM 167137).
Remarks. The extremely small size, widely open
umbilicus, and very crowded sculpture easily separate
Microcharopa mimula from all other Fijian endodont-
oids. It is most similar to the Melanesian-Polynesian
Discocharopa in size and form but obviously differs in
its spiral apical sculpture and the lack of parietal bar-
riers. The wide distribution recorded above is undoubt-
edly very incomplete because the minute size of this
species would result in its discovery by only the most
diligent collector.
Subfamily CHAROPINAE
A brief diagnosis of the Charopinae was given on
p. 70. It undoubtedly will be extensively modified when
taxa extralimital to this study are reviewed. Because
so few genera have had details of their terminal
genitalia described or figured, I have not attempted to
split the Charopinae into smaller clusters of genera. I
can emphasize that eventual generic clusters will in-
clude taxa from highly diverse geographic areas and
will deny many traditional associations. For example,
the basic anatomical structures of Notodiscus Thiele,
1931, found on Kerguelen and other Subantarctic is-
lands (Solem, 1968c), are very close to those seen in
Phenacohelix Suter, 1892, from New Zealand (fig. 11);
the structures seen in Graeffedon from Samoa (fig. 87)
are matched closely by those found in Helenoconcha
relicta Solem, 1977, from St. Helena (Solem, 1977c) and
differ markedly from any Austro-Zelandic taxa dis-
sected to date; Amphidoxa Albers, 1850, from Juan
Fernandez (fig. 30a-e) shows a combination of basic
Charopinae and unusual features and is quite unlike
the anatomical pattern of Stephanoda Albers, 1860
(fig. 31a-i), from Chile, which was treated as a sub-
genus of Amphidoxa by Pilsbry (1893-1895, pp.
39-41), Thiele (1931, p. 575), and Zilch (1959-1960, pp.
221-222). The last commentator on these two genera,
Odhner (1922, pp. 230-233, figs. 8-13, pi. 9, figs.
33-35), described the jaw and radulae of several Juan
Fernandez species, plus an external view of the
genitalia in "Stephanoda quadrata." Structures of the
latter agree quite closely with the data presented here
on Amphidoxa marmorella (Pfeiffer, 1845). Probably
the Juan Fernandez "Amphidoxa and Stephanoda" of
Odhner (1922) are monophyletic and best lumped
under Amphidoxa. They are quite distinct from the
Chilean Stephanoda. Given the above diversity and
uncertainty, no firm allocations into clusters are pos-
sible.
Fortunately, the Pacific Island taxa, with three
exceptions, appear to be a monophyletic unit. Dis-
cocharopa Iredale, 1913, with a Philippine to Society
Island range, Lagivala, known from Indonesia to Fiji
and the Ellice Islands, and the endemic Graeffedon
from Samoa and Tonga show major differences from
the other Charopinae. The remaining genera, insofar
as they have been dissected, belong to the same basic
stock as the New Zealand Charopa Albers, 1860, and
presumably will remain Charopinae regardless of fu-
ture revisions.
Discocharopa differs in shell microsculpture (figs.
5a-f, 37d). The only partial knowledge of its genital
anatomy (fig. 34a-d) makes relating it to other taxa
difficult. The extent to which the minute shell size
(mean diameter 1.36 mm.) has resulted in slenderiza-
tion and/or simplification of the anatomy is unknown.
As discussed below (p. 75), a degree of similarity to the
structures of Phenacohelix pilula (Reeve) (fig. lla-e)
exists, but their degree of relationship is unknown.
Lagivala has not been dissected and has an exten-
sive extralimital range, with species known from In-
donesia, New Guinea, and the Bismarck Archipelago.
The extent of its relationship to the Philippine-
Indonesian problematic genus Beilania Preston, 1913,
is uncertain (see p. 184). Whether Lagivala is conver-
gent with the New Zealand Mylesia (Climo, 1978) and
Ptychodon Ancey, 1888, will require detailed compari-
sons that are beyond the scope of this report. It is only
SYSTEMATIC REVIEW
73
possible to state that in barrier structure, distribution,
and basic shell features, Lagivala is not similar to
other Pacific Island genera.
Graeffedon differs quite dramatically in its anat-
omy (p. 200) and has very different barrier structure (p.
17). It is clearly related to the St. Helena endemic,
Helenoconcha, and probably will receive eventual sub-
family separation.
Of the remaining Pacific Island Charopinae,
adequate anatomical data is available for Sinployea,
Ba, and Tuimalila, fragmentary data for one species of
Vatusila, and no data on the anatomy of the monotypic
Maafu and Lauopa. The anatomical features of the
Pacific Island genera agree with those of Champa (figs.
9a-g, 10) in the basic pallial structure (except for the
lack of a glandular extension onto the pallial roof) and
the components of the interior terminal genitalia. The
penes have a verge, pocket stimulator, and circular
ridges. The lower female tracts have complex valvular
arrangements at the free oviduct and longitudinal
pilasters (compare figs. 9d and 57f). As pointed out
above (pp. 27-29), the relative prominence of these pe-
nial complex structures differs dramatically from
species to species under conditions of sympatry, but the
presence of the same exact structural elements in
Charopa, Sinployea, Tuimalila, Ba, and such extralim-
ital taxa as Amphidoxa (fig. 30e), Pilsbrycharopa (see
Solem 1970a, fig. 2i), and Pseudocharopa (fig. 32b)
indicate that this is a basic pattern. The lengthened
terminal female organs of Charopa (fig. 9b) are a minor
change of little or no phyletic significance. Charopa
(fig. 10) differs from Pacific Island taxa most noticeably
in the comparatively simple entrance of the vas def-
erens into the epiphallus. This contrasts with the
elaborate valve seen in Sinployea (fig. 57e). Despite the
extreme compaction of its genital system, the New Zea-
land Flammulina shows the same basic internal ele-
ments of the penis (fig. 29e) and has a complex vas
deferens-epiphallus junction (fig. 29i-j) equivalent to
that seen in the Pacific Island genera.
Without added reference dissections of extralimi-
tal taxa, attempts to propose a phylogenetic tree for the
Charopinae are premature. The important point here
is that many Pacific Island genera are clusterable with
New Zealand, New Guinea, Lord Howe Island, and
Juan Fernandez genera. That their degree of relation-
ship and phylogenetic order cannot yet be ascertained
is a secondary problem.
Just in reference to the Pacific Island taxa grouped
here, Sinployea is most generalized, with Ba an obvi-
ous specialization from an ancestor very similar to if
not actually the extant species Sinployea irregularis
(Garrett). The Tongan Tuimalila is an experiment in
increased whorl width and gigantism also derivable
from a Sinployea ancestor. Both Maafu and Lauopa,
neither of which have been dissected, present prob-
lems. Maafu shows great conchological specializations,
but can be derived from the Sinployea inermis groups
found in the same area. Its changes are of the same
order of magnitude found in the Lau Archipelago En-
dodontidae radiation of Zyzzyxdonta, Priceconcha, and
Thaumatodon (Solem, 1976b, pp. 461-467), and this is
interpreted as a local experiment. Lauopa, known from
limited subfossil material, is grouped here for conveni-
ence but conceivably could be an outlier of the Semper-
doninae (see p. 177). The data needed to determine its
affinities are not available. Vatusila has only a frag-
ment of its anatomy known, but the penis structures
(fig. 84a-b) are matched almost exactly by those seen
in Sinployea aunuuana (fig. 53c). I am confident that
Vatusila is correctly associated. Similarities in barrier
size, shape, and positioning (figs. 82-83, 85) to some of
the New Zealand Ptychodon andFectola (Climo, 1978,
figs. 3-4, 7-8) are intriguing, but deciding whether
this is convergence, parallelism, or genealogy is be-
yond the scope of this review. Within Vatusila, there is
a trend toward barrier reduction (see p. 192) and wider
spacing of the radial sculpture. Although it would be
tempting to suggest because of this that Vatusila
might represent the ancestor of Sinployea, I suspect
that they do not have a direct ancestor-descendant re-
lationship but represent two independent colonizations
of the Pacific Islands.
Thus, of the Charopinae on the Pacific Islands, I
interpret Ba, Tuimalila, and Maafu as local direct der-
ivations from Sinployea, with Vatusila an independent
colonization. Lauopa is of uncertain status, whereas
Discocharopa, Lagivala, and Graeffedon also represent
independent colonizations of the Pacific Islands. Be-
cause there are several independent colonizations
where external affinities are unknown, preparation of
phylogenetic trees equivalent to those of the Endodon-
tidae (Solem, 1976b, pp. 110-111, figs. 57-58) was not
attempted.
The fossil evidence is meager but intriguing. Vat-
usila eniwetokensis (Ladd, 1958) is a Miocene species
from Eniwetok in the Marshall Islands, whereas ex-
tant species are found in Lau Archipelago, Tonga,
Niue, and Vaitupu, Ellice Islands. Lagivala davidi
(Ladd, 1968) is a Late Pleistocene or Recent fossil from
Funafuti, Ellice Islands. Extant species are known
from Indonesia, New Guinea, New Britain, and Viti
Levu. The genus has not been recorded from the Sol-
omon Islands, New Hebrides, or New Caledonia. The
Funafuti species is very closely related to the Fijian L.
vivus. The Marshall Islands' Miocene Vatusila is
matched by the Miocene Cookeconcha subpacificus
(Ladd, 1958) and the Pliocene to Pleistocene Minidonta
inexpectans (Ladd, 1958) from Bikini, Marshall Is-
lands. All three genera show a shrunken post-Miocene
range correlating with the changing of the Marshall
Islands from high islands to atolls (see Solem, 1976b,
pp. 117-118). The distributional shift is thus inter-
preted as range contraction, rather than the more
simplistic assumption of migration.
Better understanding of the Pacific Island
Charopinae relationships requires more intensive in-
vestigation of the Lau Archipelago, especially to obtain
74
SOLEM: ENDODONTOID LAND SNAILS
anatomical material of Lauopa and Maafu, combined
with analysis of extralimital genera.
The generic ordering used below places two of the
"convenience-grouped" genera first (Discocharopa) and
last (Graeffedon), with Lagivala sited for comparative
purposes next to Vatusila. Detailed discussions of vari-
ation within and among genera are grouped under the
appropriate generic discussions rather than here.
Genus Discocharopa Iredale, 1913
Proc. Malacol. Soc. London, 10 (6), pp. 379-380.
Shell minute, with slightly more than 3'/2 normally coiled
whorls. Apex and spire usually slightly to moderately elevated, body
whorl descending more rapidly. Apical sculpture of major radial ribs
more crowded near end of apex, with periostracal, relatively regular
folds providing a microspiral element. Postnuclear whorls with nar-
row, sharply defined, crowded, strongly protractively sinuated radial
ribs. Microsculpture of very fine radial riblets with serrated edges,
no trace of spiral sculpture. Umbilicus very widely open, saucer-
shaped, regularly decoiling, margins rounded. Whorls flattened lat-
erally above and below rounded periphery. Color white with a yellow
tinge. Aperture with or without a small to very large and deeply
recessed, medial parietal barrier. Pallial region with complete sec-
ondary ureter, kidney bilobed, pericardial lobe longer than rectal.
Genitalia with short bilobed ovotestis, slender talon. Terminal
genitalia slender with subapical entrance of vas deferens, no exter-
nally recognizable epiphallus, penis internally with longitudinal
pilasters.
Type species. Charopa (Discocharopa) exquisita
Iredale, 1913, by original designation.
The shell sculpture of Discocharopa (figs. 5a-f,
37d) is quite different from that found in any other
Pacific Island Charopidae. Like the Endodontidae, the
apical whorls (fig. 5b-c) have prominent radial ribs
that become quite crowded near the nuclear-post-
nuclear boundary (fig. 5a), but the microsculpture is
quite different from that seen in the Endodontidae. The
latter have very fine, regular, "squiggly" microspirals
that continue onto at least the early postnuclear
whorls (Solem, 1976b, p. 38, fig. 28c-e; p. 39, fig. 29a).
The Charopidae have fine to prominent spiral cords
(for example Sinployea modicella, fig la-c, p. 10) with
weak to strong (fig. 2c, p. 11) secondary radial ele-
ments. Discocharopa has spiral sculpture that is of ir-
regular shape, length, spt cing, and angle of orienta-
tion. Similar apical sculpture has been found in several
New Zealand species usually classified in Ptychodon
(for example, P. microundulata in Solem, 1970b, pi. 58,
figs. 4-6) but now referred to Mylesia (see Climo, 1978,
p. 185). The postnuclear sculpture in Discocharopa
differs from all endodontoids examined to date. All
these have both microspiral and microradial elements,
but Discocharopa (fig. 5b, d-f) shows no trace of micro-
spiral elements, and the radial sculpture has unusual
features. The major ribs (fig. 5a-b, d-f) retain rem-
nants of microradial clustering on their apices only on
early postnuclear whorls (a-b), whereas the latter
whorls (d-e) appear to have a "rolled" or expanded rib
peak. The microradials (e-f) are partly broken up into
irregular short segments by transverse grooves. The
latter have no counterpart in any Pacific Island species
examined to date.
After considerable effort over many years, it was
possible to obtain material and partially work out the
basic anatomy of Discocharopa. The wide range of Dis-
cocharopa yielded hope of borrowing material to dis-
sect, and fieldwork by L. Price in Fiji and by Price and
Solem in Australia could have produced live material.
No material was obtained until two specimens from the
National Museum, New Zealand, were provided for
study by their collector, Frank Climo. They were col-
lected "behind the landing," Raoul Island, Kermadec
Islands, on August 24, 1972, and thus represent near
topotypic material of the genotype, Discocharopa ex-
quisita (Iredale, 1913, pp. 379-380), which was de-
scribed from "Sunday Island, Kermadec Group. Living
under rotten wood, stones, etc."
Both examples had completely retracted within
their shells, and the albumen gland had crystallized,
which made interpretation of most structures difficult.
The main uncertainties include whether the prostate-
uterus are combined or separated, the talon structure,
and the position and shape of the spermathecal head. I
suspect that the latter is bound into the aorta at the
area where the kidney base and albumen gland touch
the diaphragm. In neither specimen was I successful in
separating the kidney base, albumen gland, intestines,
and hindgut because of extensive albumen gland crys-
tallization. Undoubtedly the spermathecal head was
mangled and lost in the attempts to work out
structures in this area. The exact shape, size, and re-
lationship of the talon to the hermaphroditic duct could
not be determined precisely. Figure 34b represents the
best interpretation of dissector and illustrator, but
could be incorrect.
The ovotestis (fig. 34a, G) is unusual in that it lay
alongside the stomach and below the main area of di-
gestive gland. The bilobed condition is found in many
charopids, but the anterior shift in position has not
been seen by me in other taxa. The hermaphroditic
duct (GD) was iridescent, again as in many charopids,
and the albumen gland was deeply dented by loops of
the intestine, thus having a very irregular shape. A
swollen uterine area was clear, but the prostate tissue
was very small and the spermathecal shaft closely
bound to the margin. Because of partial crystallization
in the uterine area, details could not be worked out.
The vas deferens broke and is shown (VD) reflexed, but
the long vagina and penis have been unfolded in order
to show a little more detail (fig. 34c). The entrance of
the vas deferens into the penis is subterminal, with a
short penial retractor muscle inserted apically. Appar-
ently there are at least two longitudinal pilasters
within the penis, but exact structures are too small to
be determined from available material.
The pallial region (fig. 34d) has a closed and com-
plete secondary ureter (KD); the kidney (K) is bilobed
with a large section touching to partly overlapping the
SYSTEMATIC REVIEW
75
GG
FIG. 34. Anatomy of Discocharopa aperta (MbllendorfD. Forest
behind landing, Raoul Id., Kermadec Islands. F. Climo! VIII-24-1972.
FMNH 193764: a, ovotestis; b, detail of talon and carrefour; c, post-
apical genitalia; d, pallial region. Scale lines as marked. (EL).
hindgut (HG). Details of the anus (A) and external
ureteric pore (KX) are only approximate. The apex of
the kidney is curved over around the start of the ure-
ter, possibly as a result of retraction into the shell.
The larger of the two dissected specimens was 0.62
mm. high, 1.51 mm. in diameter, H/D ratio 0.413,
whorls 3%, umbilicus 0.43 mm. wide, D/U ratio 3.54.
The more successful dissection was made on the
smaller example, which was 0.58 mm. high, 1.25 mm.
in diameter, with 3% whorls.
All observed anatomical features are consistent
with Discocharopa being a member of the Charopinae.
Unfortunately the small size, contraction, and partial
crystallization combined to prevent full study.
Whether the slender terminal genitalia is a secondary
specialization is uncertain. Unquestionably the fea-
tures observed are different from Sinployea and its re-
latives. To a certain extent there are similarities with
Phenacohelix (fig. lie, e) in general penis shape,
ovotestis, and pallial configuration, but there is a clear
difference in vas deferens insertion, talon, and termi-
nal female genitalia configurations. Of the extralimi-
tal taxa considered here, Discocharopa has the most
similarities to Phenacohelix, but the degree of their
relationship remains unknown.
Because only partial anatomical data was re-
covered, this general discussion substitutes for a for-
mal description.
Originally, only the Kermadec Islands populations
and the Tasmanian D. bassi (Legrand, 1871) were
placed in Discocharopa. Subsequently, Iredale (1937a,
p. 325) included several additional Australian and
Tasmanian species, Cotton (1939, p. 176) described a
Paralaoma as belonging to this genus, van Benthem
Jutting (1951) named/), microdiscus from the Celebes,
and I (Solem, 1957, 1959a) added Philippine and New
Hebridean (fig. 36) taxa. None of the Tasmanian
species, D. vigens (Legrand, 1871), D. bassi (Legrand,
1871), D. lottah (Petterd, 1879), orD. mimosa (Petterd,
1879), appear to be correctly classified in Discocharopa,
but generic reallocation is not attempted here. Dis-
cocharopa concinna (Hedley, 1901) and D. planor-
bulina (Tate, 1896) from Queensland and the Northern
Territory of Australia are synonymized with D. aperta
(Mollendorff) from the Philippines. The latter is the
oldest name and applies to the single species recog-
nized. Discocharopa werneri Solem (1957, pp. 4-6, fig.
2) from Mindanao is totally unrelated to Discocharopa.
Its original assignment here was a desperation effort to
avoid description of a new genus, compounded by an
unwillingness to place it in a New Zealand genus when
adequate comparative material was lacking.
Combining all described species with D. aperta is
based upon examination of all types and study of over
300 specimens. This results in recognizing a species
with the widest distribution of any endodontoid, ex-
tending from the northern Philippines and Java to
Central Australia, Kermadec Islands, and the Society
Islands. The only approximately equivalent distribu-
tion is that shown by Stenopylis coarctata (Mollendorff,
1894), which has almost the same Indonesian and
Australian range, but only reaches the Solomon Is-
lands and has not been found in the New Hebrides, Fiji,
Kermadec Islands, or Polynesia. Data on this species
are summarized by Solem (1957, pp. 8-11, fig. 4). Sub-
sequently it was reported from Bach Long Vi (Nightin-
gale Island) in the Gulf of Tonkin (Saurin, 1960, pp.
7-9, fig. 3, pi. 1; figs. 6a-b, 7) and was mistakenly
described as a new species, Microphyura nightingali.
Microphyura is a New Caledonian endemic rhytidid
genus whose conchological appearance is convergent
with that of Stenopylis. Hedley (in Tate, 1896, pp.
221-222, fig. C) illustrated the jaw and radula of
Stenopylis. Tentatively, I (Solem, 1975) classified it in
the Helicodiscidae, an otherwise North American com-
plex, and subsequent dissection has confirmed this as-
signment (Solem, unpublished data).
Discocharopa and Stenopylis have been collected
together on several occasions, and I suspect their
ecological requirements are very similar. The method
of apertural narrowing is very different, with Dis-
cocharopa using a single, deeply recessed parietal,
whereas Stenopylis uses a combination of lip thicken-
76
SOLEM: ENDODONTOID LAND SNAILS
ing and internal tubercles (Solem, 1957, p. 11, fig.
4b-c, e).
Differences of Discocharopa from other genera are
gross in nature. Microcharopa from Fiji is most apt to
be confused in size but is slightly smaller (mean diame-
ter 1.07 mm.), has strong spiral apical sculpture (fig.
4a-e, p. 13), proportionately larger apical whorls, and
more evenly rounded body whorl contours. Punctum
has a much higher spire, is distinctly smaller, has
prominent spiral apical cords and a narrower um-
bilicus. No Pacific Island Charopidae without promi-
nent apertural barriers average less than 2.00 mm. in
diameter, whereas those Lagiuala and Vatusila in the
1.40-1.75 mm. size range have very prominent aper-
tural barriers at or very near the lip edge. Most of these
have marked lateral body whorl compression. Of the
six Endodontidae averaging less than 2 mm. in diame-
ter, only the Manu'a Group Minidonta manuaenis
Solem 1976, occurs within the geographic range of Dis-
cocharopa. That species has more than four whorls, a
very narrow umbilicus, H/D ratio over 0.550, and seven
prominent apertural barriers. The other species of
Minidonta are Hawaiian, Mangarevan, and the Bikini
Atoll fossil, M. inexpectans (Ladd, 1958).
Discocharopa aperta (Mollendorff, 1888). Figures
5a-f, 34a-d, 35a-b, 36a-c, 37a-d.
Patula aperta Mollendorff, 1888, Nachr. Bl. deut. Malak. Gesell.,
20 (5-6), p. 89 Montalban, Rizal, Luzon, Philippines; Pilsbry,
1892, Man. Conchol., (2) 8, pp. 80-81, pi. 37, figs. 35-37; Mol-
lendorff, 1898, Abhl. Naturf. Ges. Gorlitz, 22, p. 88 Luzon,
Leyte, Calamianes.
Endodonta (Champa) planorbulina Tate, 1896, Rep. Horn Sci.
Exped. Central Australia, (2), p. 187, pi. 17, fig. 3 Palm
Creek, Krichauff Range, Northern Territories, Australia.
Endodonta concinna Hedley, 1901, Proc. Linn. Soc. New South
Wales, 25, p. 729, pi. 48, figs. 1-3 Bundaberg, Queensland,
Australia.
Charopa (Discocharopa) exquisita Iredale, 1913, Proc. Malacol.
Soc. London, 10 (6), pp. 379-380, pi. XVIII, fig. 8 Sunday
Island, Kermadec Group.
Ptychodon celebica B. Rensch, 1932 (not Sarasin & Sarasin, 1899),
Zool. Jahrb., Syst, 63, p. 101 Sumba; B. Rensch, 1935, Sitz.-
Ber. Gesell. Naturf. Freunde, Berlin, 1935, p. 322 Timor.
Pyramidula aperta (Mollendorff), Faustino, 1930, Philippine J.
Sci. ,42(1), p. 110.
Discocharopa planorbulina (Tate), Iredale, 1937, Australian Zool.,
8 (4), p. 325; Iredale, 1937, South Australian Nat., 18 (2), p. 25,
pi. 1, fig. 21.
Discocharopa concinna (Hedley), Iredale, 1937, Australian Zool., 8
(4), p. 325.
Charopa exquisita (Iredale), I. Rensch, 1937, Arch. f. Naturgesch.,
n. f., 6 (4), pp. 590-591 Malkong-Bach, Weiten Buch and
Vunapope, Gazelle Peninsula, New Britain, Bismarck Ar-
chipelago.
Charopa (Discocharopa) microdiscus van Benthem Jutting, 1951,
Basteria, 15 (1-2), pp. 28-29, fig. 1 South Celebes and West
Java; van Benthem Jutting, 1952, Treubia, 21 (2), p. 398
West Java; van Benthem Jutting, 1953, ibid., 22 (2), p. 302
Ambon.
Discocharopa aperta (Mollendorff), Solem, 1957, Fieldiana: Zool-
ogy, 42 (1), pp. 3-4, fig. 1, a-d Luzon and Catanduanes,
Philippine Islands.
Discocharopa microdiscus van Benthem Jutting, Solem, 1958,
Arch. f. Mollusk., 87 (1-3), p. 21.
FIG. 35. Parietal barrier variation in Discocharopa aperta (Mol-
lendorff): a, Tjamba Pass, near Makassar, Celebes. ZMA: b, Blue
Lake, Sunday Island, Kermadec Islands. Paratype of Charopa ex-
quisita Iredale, 1913. FMNH 117214. Scale line equals 1 mm. (JC).
Charopa microdiscus van Benthem Jutting, 1958, Verh. Naturf.
Ges. Basel, 69 (1), p. 106 Sumba, Indonesia.
Discocharopa planulata Solem, 1959, Fieldiana: Zoology, 43 ( 1), pp.
82-83, pi. 32, figs. 1-3 stream drift in Sarakata River Valley,
Espiritu Santo, New Hebrides.
Diagnosis. Shell minute, diameter 1.18-1.84 mm. (mean 1.36
mm.), with 3*78-4(4 normally coiled whorls. Apex and spire rarely
flat, usually slightly to moderately elevated, body whorl descending
more rapidly to very strongly deflected (Rurutu, Austral Islands),
spire protrusion 'Az to more than 4 body whorl width, usually about
l k, H/D ratio 0.341-0.615 2 (mean 0.438). Apical sculpture of radial
ribs that on lower portion of apex becomes more crowded, with barely
visible spirals. Postnuclear whorls with prominent, narrow, sharply
defined, strongly protractively sinuated radial ribs, 77-148 (mean
110.6) on the body whorl, whose interstices usually are 2-4 times
their width. Ribs/mm. 18.0-40.4 (mean 26.9). Microsculpture of very
fine radial riblets, 3-5 between each pair of major ribs, no secondary
spiral sculpture. Umbilicus very widely open, saucer-shaped, regu-
larly decoiling, contained 2.00-3.21 times (mean 2.58) in the diame-
ter, margins rounded. Sutures impressed, whorls strongly rounded
above and on basal margin, usually very slightly to slightly com-
pressed laterally above and below evenly rounded periphery. Aper-
ture subcircular, moderately to strongly compressed laterally above
periphery, less strongly compressed laterally below periphery, in-
clined 10-30 from shell axis. Parietal wall with or without a medial
barrier, which, when present, is usually recessed Vt to more than !4
whorl and varies from a low cordlike ridge extending 3 Ae of a whorl to
a very high crescentic lamella of up to % aperture height, with
gradual anterior and posterior descensions, moderately expanded
and weakly serrated medially when very high. Presence or absence
on either a populational or individual basis. Color white, with at
most a faint yellow tinge.
In having stronger radial than spiral apical
sculpture, a very widely open umbilicus, and at most
only a single very deeply recessed parietal barrier, Dis-
cocharopa aperta is immediately differentiated from
other species of similar size. Species of Beilania and
2 The only two specimens with H/D ratio greater than 0.500
(0.549, 0.615) are Rurutu examples with greatly deflected body
whorls.
SYSTEMATIC REVIEW
77
Lagivala have spiral apical sculpture predominating,
more apertural barriers that are near the lip edge (at
least for the parietals), and generally much thicker
body whorls with less sinuated sculpture. Mi-
crocharopa mimula is most similar in size and shape,
but differs in its very distinctive apical sculpture (fig.
4), lower whorl count, lack of apertural barriers, and
proportionately much larger apical whorls.
Description of Patula aperta. Shell minute, with slightly less
than 3% normally coiled whorls. Apex flat, spire slightly elevated,
body whorl descending a little, H/D ratio 0.443. Apical whorls l'/2,
sculpture partially eroded, but traces of radial ribbing remaining.
Postnuclear whorls with narrow, somewhat irregular radial ribs,
eroded over much of the shell. Microsculpture obscured by incrusta-
tions and eroding of surface except for faint traces in umbilicus.
Sutures deep, whorls strongly rounded above, somewhat flattened
laterally above periphery with evenly rounded outer margins. Um-
bilicus saucer-shaped, regularly decoiling, contained 2.51 times in
the diameter, with evenly rounded margins. Color white, with
periostracal remnants having a faint yellow tone. Aperture subcircu-
lar, slightly flattened laterally above periphery, inclined about 15
from the shell axis. No parietal barrier visible from aperture. Height
of lectotype 0.64 mm., diameter 1.45 mm.
Lectotype. Philippine Islands: Luzon, Montalban.
SMF 165358.
Description of Endodonta (Charopa) planorbulina. Shell mi-
nute, with slightly more than 3Vi normally coiled whorls. Apex and
spire barely protruding, last whorl not descending, H/D ratio 0.400.
Apical whorls 1%, surface badly eroded with only faint traces of
regularly spaced radial ribbing remaining. Postnuclear whorls with
moderately widely spaced, protractively sinuated radial ribs, 91 on
the body whorl, whose interstices are 2-3 times their width. Micro-
sculpture mainly eroded, consisting, when visible, of faint radials.
Sutures relatively deep, whorls strongly rounded above, slightly flat-
tened laterally above periphery with evenly rounded outer margin.
Color white. Umbilicus saucer-shaped, regularly decoiling, con-
tained 2.70 times in the diameter with rounded margins. Aperture
subcircular, slightly flattened laterally above periphery, inclined
about 10 from shell axis. Height of holotype 0.61 mm., diameter 1.52
mm.
Holotype. Australia: Palm Creek, Krichauff
Range, Northern Territories. SAM D.3222.
Description of Endodonta concinna. Shell minute, with 3'/a rel-
atively loosely coiled whorls. Apex and spire barely elevated, last
half of body whorl descending moderately, H/D ratio 0.445. Apical
whorls 1%, sculpture of relatively widely spaced radial ribs with an
extremely fine microsculpture of more crowded spiral riblets. Re-
maining whorls with narrow, relatively crowded, protractively
sinuated radial ribs, about 102 on the body whorl, whose interstices
are 2-4 times their width. Microsculpture barely visible, consisting
of fine radial riblets. Sutures relatively impressed, whorls strongly
rounded above, flattened laterally above periphery and partially on
basal margin. Color very light yellowish white. Umbilicus saucer-
shaped, regularly decoiling, contained 2.81 times in the diameter.
Aperture ovate, compressed laterally above periphery and on basal
margin, inclined about 15 from the shell axis. Height of lectotype
0.61 mm., diameter 1.48 mm.
Lectotype. Queensland: Bundaberg. AMS C.8970.
Description of Charopa (Discocharopa)exquisita. Shell minute,
with 314 normally coiled whorls that only slightly increase in width.
Apex and spire barely emergent, body whorl descending slightly,
H/D ratio 0.450. Apical whorls l'/2, first quarter whorl worn smooth,
lower sculpture of prominent, narrow radial ribs, more widely spaced
above, becoming crowded near end, with microsculpture of very fine
spiral ribbing. Remaining whorls with narrow, strongly protrac-
tively sinuated lamellar radial ribs, 100 on the body whorl, whose
interstices are 3-5 times their width. Microsculpture of very faint
radial riblets. Sutures deeply impressed, whorls evenly rounded.
Umbilicus saucer-shaped, broadly opened, regularly decoiling, con-
tained 2.58 times in the diameter. Color light yellow-horn without
darker maculations. Aperture circular, flattened laterally above
periphery, inclined about 20 from shell axis. Parietal wall with
single, very deeply recessed barrier visible through shell. Height of
lectotype 0.59 mm., diameter 1.32 mm.
Lectotype. Kermadec Islands: crater slopes
northeast of Blue Lake, Sunday Island. Collected
under stones by W. R. B. Oliver on October 3, 1908.
BMW.
Description of Charopa (Discocharopa) microdiscus. Shell
minute, with slightly more than 3% normally coiled whorls. Apex
and spire slightly and evenly elevated, body whorl descending a little
more rapidly, H/D ratio 0.424. Apical whorls 1%, sculpture of nar-
row, prominent, moderately widely spaced radial ribs, becoming
more crowded near end with a much finer, very inconspicuous mi-
crosculpture of spiral ribs. Postnuclear whorls with fine, strongly
protractively sinuated lamellar radial ribs, 111 on the body whorl,
whose interstices are 2-3 times their width. Microsculpture barely
visible under extreme magnification, consisting of very fine radial
riblets. Color very faint yellowish white. Sutures deep, whorls
strongly rounded above, flattened laterally above and below evenly
rounded periphery with slightly flattened columellar margin. Um-
bilicus very broadly open, saucer-shaped, regularly decoiling, con-
tained 2.25 times in the diameter. Aperture subcircular, flattened
laterally above and below periphery, inclined about 15 from the
shell axis. No parietal barrier visible from aperture. Height of
holotype 0.69 mm., diameter 1.63 mm.
Holotype. South Celebes: hills of Pangkadjene,
Makassar. ZMA.
Range. Philippines and Java east through
Sumba, Timor, and Biak to the Bismarck Archipelago,
New Hebrides, and Fiji, Kermadec Islands, Manu'a
Group in Samoa, Rurutu in the Austral Islands,
Borabora in the Society Islands, Northern Australia as
far south as the MacDonnell Ranges and Northern Vic-
toria, and Bundaberg in coastal Queensland. Probably
in most intermediate areas.
Material. Philippine Islands: north Luzon (4
specimens, RSME); Montalban (7 specimens SMF
165358-60, NMWC). Catanduanes (1 specimen,
FMNH 57193). Panglao (1 specimen, RSM). Cala-
mianes: Koron (3 specimens, SMF 165361). Mindanao:
Samial (5 specimens, SMF 165362). Busuanga (4
specimens, SMF 165363, FMNH 18725). Bulacan:
Bayabac (7 specimens, SMF 165364-5).
Java: Koeripan, Buitzenborg (2 specimens, ZMA,
collected February 21, 1932, by van Benthem Jutting).
Sumba: Waikarudi (2 specimens, NHB 6092-c);
Langgaliru (1 specimen, NHB 6092-d); Kedi (1 speci-
men, NHB 6092-b); Mau Marru (5 specimens, NHB
6092-a, ZMA).
Ambon (2 specimens, ZMA, collected October
11-13, 1949, by M. A. Lieftinck).
South Celebes: Pangkadjene near Makassar (21
specimens, ZMA, collected May 9 and October 14, 1948,
by G. A. Tammes-Bolt); Palopo near Rante Pao, 23 km.
from Makale at 700-800 m. elevation (1 specimen,
ZMA, collected October 13, 1948, by G. A. Tammes-
Bolt); Kalkrotsenlangs Grote, on way from Makale,
78
SOLEM: ENDODONTOID LAND SNAILS
near Kalossi at 700-800 m. elevation (4 specimens,
ZMA, collected September, 1948, by G. A. Tammes-
Bolt).
Timor: Araki-Vlakte (4 specimens, ZMA, collected
April 12, 1950, by B. Polak); Niki-Niki (5 specimens,
NHB 4169-a); Noilmina (40 specimens, ZMB, FMNH
146025).
Biak: Zuid-Biak Subdistrict (Station G-24) (olim
Sorido) (1 specimen, ZMA, collected March 26, 1952).
Aru Islands: Wokan (1 specimen, MSNG).
Queensland: Bundaberg (9 specimens, AMS
C.8970, FMNH 117331).
Northern Territory: Palm Creek, Krichauff Range,
near Hermannsburg (3 specimens, SAM D.3222); Sta-
tion WA-130, Cycad Gorge, Palm Creek, Krichauff
Range (7 specimens, FMNH 171558); Station WA-113,
Glen Helen entrance to Finke Gorge, MacDonnell
Ranges (1 specimen, FMNH 182104); Station WA-133,
Temple Bar Gap, MacDonnell Ranges (1 specimen,
FMNH 182119); Station WA-446, Kathleen Spring,
Gill Range (3 specimens, FMNH 200476).
Western Australia: Stations WA-303 and WA-305,
0.5-1.0 km. east of Yammera Gap, Napier Range (2
specimens, FMNH 199179, FMNH 199245); northwest
of Stumpy's Well, southwest side Napier Range (1
specimen, WAM 824-76); "The Tunnel," Tunnel
Creek, Napier Range (1 specimen, WAM 1758-78);
Stations WA-257 and WA-258, Brooking Gorge, Oscar
Ranges (9 specimens, FMNH 199491, FMNH 199547);
Station WA-255, near Gieke Gorge Ranger station,
Gieke Gorge National Park, Oscar Ranges (3 speci-
mens, FMNH 199597); Cave Spring, Bugle Gap, Law-
ford Ranges (WAM).
New Britain: Vunapope (12 specimens, ZMB,
FMNH 146027); Malkong (3 specimens, ZMB).
New Hebrides: Vila, Vate (3 specimens, BMNH
1915.12.31.602, AMS C. 15679); Sarakata River, Es-
piritu Santo (3 specimens, MCZ 186826, FMNH 54904,
BPBM 212378).
Kermadec Islands: Sunday Island (34 specimens,
AIM, IRB, DMW 12449); under pumice on Mt. Junction
(4 specimens, DMW 12424, collected September 18,
1908, by W. R. B. Oliver); under stones on crater slope,
northeast of Blue Lake (14 specimens, DMW 12423,
FMNH 117214, collected October 3, 1908, by W. R. B.
Oliver).
Fiji: Viti Levu: Sigatoka, near base of limestone
cliff, Vz-V/z miles inland (1 specimen, BPBM 87930,
collected July 12, 1928, by H. S. Ladd); Lami Ridge
(Station 47), 5 miles west of Suva at 5-150 ft. elevation
(2 specimens, BPBM 178489, collected July 24, 1938,
by Y. Kondo and C. M. Cooke, Jr.).
Fiji: Lau Archipelago: Yangasa Levu, south end
(Station 30), about 150 ft. inland at 40 ft. elevation (17
specimens, BPBM 167137, collected July 26, 1934, by
H. S. Ladd); Navutu-I-Loma (1 specimen, BPBM
167102); northeast quarter (Station 28), 150 yd. in-
land at 100 ft. elevation (3 specimens, BPBM 167017,
collected July 24, 1934, by H. S. Ladd); Namuka (1
specimen, BPBM 166668): Matandolo (Station 2),
north-central point of island, 500 ft. inland at 65-75 ft.
elevation (6 specimens, BPBM 166614, collected July
7, 1934, by H. S. Ladd); Wangava (4 specimens, BPBM
166994): northeast end (Station 27), V* mile inland at
75 ft. elevation (3 specimens, BPBM 166965, collected
July 22, 1934, by H. S. Ladd); Karoni (1 specimen,
AIM, collected July 4, 1977, by Walter Cernohorsky):
slope of peak at about 90 ft. elevation (6 specimens,
BPBM 78603, BPBM 78606, collected August 15, 1924,
by E. H. Bryan, Jr.); Aiwa: southwest quarter of island
(Station 40), 50 yd. inland at 25 ft. elevation (1 speci-
men, BPBM 167208, collected August 7, 1934, by H. S.
Ladd); Nayau: Nauko (Station 43), Vz mile inland at
base of limestone cliff, 250 ft. elevation (3 specimens,
BPBM 167230, collected August 13, 1934, by H. S.
Ladd); Vanua Mbalavu: between Valika and Mosomo
Bay (Station 78), limestone hill about % mile inland at
200-250 ft. elevation (4 specimens, BPBM 179582-3,
collected August 9, 1938, by E. C. Zimmerman and Y.
Kondo).
Samoa: Aunuu Island off Tutuila: 10 yd. east of
lighthouse, 200 yd. inland at 250 ft. elevation (4 speci-
mens, BPBM 83243, collected February 4, 1926, by T.
Dranga) (1 specimen, BPBM 171020).
Ta'u: Siufaga (Station 76) (1 specimen, BPBM
171074); Tavalagi Ridge (Station 75), Va mile inland at
450 ft. elevation (27 specimens, BPBM 171066, BPBM
171107); Lepuu Cliffs (4 specimens, BPBM 171126).
Olosega: Olosega Village (1 specimen ex BPBM
188720).
Society Islands: Borabora, south slope (Station
1093) of Pahio-Temanu ridge, 800 ft. elevation in dirt
at cave entrance (1 specimen, BPBM 152396, collected
October 13, 1934, by Gessler and St. John).
Austral Islands: Rurutu: Mato Naa, bluff north of
Moerai (Station 760), 20-30 yd. inland at 5-15 ft. ele-
vation (6 specimens, BPBM 148292); Mato Arei (Sta-
tion 775), cliff southeast of Moerai at 50-150 ft. eleva-
tion (6 specimens, BPBM 148559, collected August 27,
1934, by Y. Kondo and D. Anderson).
Remarks. Except for collections from river drift
or detritus sweepings, Discocharopa aperta (Mollen-
dorff) is represented in collections by very small sam-
ples. Partly this results from its small size, but more
probably from the secretive habits and actual rarity.
Iredale (1913, p. 366) reported that specimens were
found under "a piece of black bark," later "on the
underside of moss-covered stones deeply imbedded in
earth on the side of the cliff," a "few . . . under dead
nikau leaves," and "under loose dry pumice stones in
the crater." He reported that "Only two or three were
found at a time." No other data have been recorded
concerning the habits of living specimens.
At different times since 1961, I have been able to
examine the types of all described species. Some were
restudied a second time in order to check on details
overlooked initially or to confirm structures. Despite
SYSTEMATIC REVIEW
79
the variations outlined below, I conclude that only one
species can be recognized. My list of material covers all
the references cited above except for that of'Ptychodon
celebica" from B. Rensch (1932, p. 101; 1935, p. 322).
Broken or juvenile examples from these materials were
seen in the SMF collection, but were not measured or
listed. They are Discocharopa, however, and not
Beilania philippinensis (Semper). Description of sev-
eral species was caused by the wide geographic range,
large shape variation, and the short diagnoses origi-
nally presented. I am among the guilty, since Dis-
cocharopa planulata Solem (1959a) from the New Heb-
rides is based on rather large and widely umbilicated
shells (fig. 36a-c). Previously (Solem, 1957, p. 4) I had
suggested that the Philippine/), aperta and Indonesian
D. microdiscus might be synonymous, but failed to ex-
tend this to the other taxa.
Data on size and shape variation are summarized
in Table XIII. The growth pattern is such that certain
separation of adult from subadult examples is very dif-
ficult with fresh material and virtually impossible
with worn, dead shells from river drift or detritus. Cal-
culations have been restricted to means and ranges
under these circumstances. Despite a certain partial
mixture of subadults and adults, there is quite minor
variation in size and shape. Specimens from the Bis-
marck Archipelago are distinctly smaller and have
more numerous, crowded radial ribs. Australian
TABLE XIII. - GEOGRAPHIC VARIATION IN DISCOCHAROPA APERTA.
NUMBER OF
NAME SPECIMENS
RIBS
RIBS/MM.
HEIGHT
DIAMETER
Montalban, Luzon,
11
102
.0(99-105)
23.
7(23
,2-24,
0)
0.
58(0.49-0
.66)
1.38(1
.27-1.53)
Philippines
Northern Luzon
5
104
.7(100-110)
24.
6(24
,0-25,
4)
0.
59(0.53-0
.66)
1.31(1
.15-1.38)
Sumba and Celebes
12
25
101
100
.4(78-123)
.9(77-127)
26.
24.
3(21
4(18
,8-31,
,0-34.
3)
7)
0.
0.
55(0.42-0
56(0.48-0
.69)
.69)
1.27(0
1.34(1
.99-1.63)
.09-1.55)
Timor
Mew Britain.
11
133
.0(118-148)
35.
3(30.
,2-40.
6)
0.
65(0.48-0
.67)
1.21(1
.07-1.45)
Bismarcks
Krichauff Mts..
3
88
.3(84-91)
19.
7(19.
2-20.
8)
0.
59(0.58-0
.61)
1.43(1
.38-1.51)
N. Territories.
Australia
Bundaberg.
Queensland, Australia
5
102
21.
9
0.
63(0.59-0
.67)
1.47(1
.40-1.56)
New Hebrides
5
127
.3(125-131)
29.
5(29.
2-29.
6)
0.
54(0.46-0
.63)
1.36(1
.25-1.43)
Fill. Lau Group
7
3
15
135
140
114
.7(125-147)
.0(102-128)
31.
29.
28.
1(28,
2
0(25,
5-35,
4-30.
7)
5)
0.
0.
0.
64(0.53-0
58(0.46-0
64(0.49-0
.74)
.70)
.72)
1.56(1
1.39(1
1.32(1
.32-1.84)
.25-1.53)
.07-1.45)
Fiii. Viti Levu
Samoa
Rurutu. Austral Is.
6
1
22
109
103
.7(85-123)
25.
25.
8
1(20,
9-30.
6)
0.
0.
0.
70(0.63-0
64
57(0.49-0
.79)
.63)
1.37(1
1.43
1.32(1
.28-1.48)
.23-1.41)
Borabora. Society Is.
Sunday Id. .,
Kermadec Is.
Monta.
N. Luz.
H/D
0.418(0
0.452(0
RATIO
.385-0
.425-0
.476)
.476)
WHORLS
3 l/2-(3 1/4-3 3/4)
3 3/8(3 1/8-3 1/2)
UMBILICUS
0.56(0.53-0.65)
0.52(0.43-0.58)
n/u
2.57(2
2.53(2
RATIO
.40-2.96)
,40-2.69)
BODY WHORL
WIDTH SP/BWW
Sumba
Timor
N. Bri.
Krich.
Bunda.
N. Heb.
0.430(0
0.419(0
0.444(0
0.409(0
0.428(0
0.400(0
.396-0
. 382-0
.402-0
.400-0
.404-0
.341-0
.487)
.488)
.466)
.417)
.445)
.437)
3 l/2-(3 1/8-4)
3 1/2-O-3 7/8)
3 3/8+(3 1/4-3 5/8)
3 1/8(3-3 1/4)
3 l/2+(3 1/2-3 3/4)
3 1/2(3 1/4-3 5/8)
0.53(0.
0.53(0.
0.49(0.
0.54(0.
0.55(0.
0.60(0.
39-0.72)
39-0.61)
43-0.59)
51-0.56)
51-0.64)
53-0.66)
2.39(2
2.57(2
2.48(2
2.66(2
2.68(2,
2.29(2,
,22-2
,36-2
,38-2
,57-2
,44-2
,16-2
.63)
.77)
.58)
.71)
.81)
.63)
0.38 0.130
Lau
0.410(0
. 390-0
.450)
3 3/4+(3
1/2-4 1/4)
0.72(0.
48-0.92)
2.21(2,
00-2
.76)
0.40(0.35-0.44) 0.178(0.111-0.292)
V. Levu
0.418(0
. 368-0
.457)
3 3/8+(3
1/4-3 5/8)
0.46(0.44-0.48)
3.04(2.
81-3
.21)
Samoa
0.486(0
.429-0
.524)
3 l/2-(3
1/4-3 5/8)
0.47(0.
39-0.61)
2.80(2.
59-3
.09)
0.39(0.34-0.41 0.171(0.091-0.286)
Rurutu
0.514(0
.444-0
.615)
3 5/8(3
3/8-4)
0.51(0.
41-0.59)
2.71(2.
50-3
.12)
0.41(0.38-0.44) 0.208(0.146-0.240)
Borab.
0.448
3 1/2+
0.61
2.35
0.41 0.120
Sunday
0.434(0
. 390-0
.469)
3 l/4+(3-3 1/2)
0.48(0.
46-0.56)
2.57(2.
42-2
.68)
80
SOLEM: ENDODONTOID LAND SNAILS
*** ^^
FIG. 36. a-c, Discocharopa aperta (Mollendorff) from the New
Hebrides. River drift, Sarakata River, Espiritu Santo, New Hebrides.
Holotype of Discocharopa planulata Solem, 1959. UMMZ 186037.
Scale line equals 1 mm. (Solem, 1959a, pi. 32, figs. 1-3). (EJP).
specimens are large and have fewer and more widely
spaced radial ribs (fig. 37a-d). Lau Archipelago shells
are very large and widely umbilicated, with the New
Hebridean shells only slightly smaller (fig. 36a-c).
Both have intermediate ribbing frequency and spacing.
Shells from Viti Levu and Samoa are much more nar-
rowly umbilicated. The higher H/D ratio and narrower
umbilicus in the Rurutu shells results from drastic de-
flection of the body whorl. Otherwise, differences are
well within the range of statistical error.
Probably more importance can be attached to var-
iation in the parietal barrier. Kermadec shells (fig.
35b) have a very large barrier recessed about one-
quarter whorl and usually visible through the shell,
but not from the aperture. Rurutu examples have a
slightly smaller parietal that was clearly visible from
the aperture, as did the single example from the Soci-
ety Islands. Specimens from the Lau Archipelago had a
parietal visible in juveniles, sometimes large and
sometimes small, but not visible from the aperture in
any adults. No Lau Archipelago adults had a translu-
cent shell, but unless there was deeper recession of the
barrier in adults, it should have been visible. No
Australian, Samoan, Viti Levu, or New Hebridean
shells showed any trace of a parietal.
A number of Samoan shells were sufficiently
translucent to allow confirmation of the parietal's ab-
sence. Some Philippine specimens had a low to medium
parietal, but most showed no trace of a barrier. Most
Sumba examples had a large parietal, whereas the
Celebes shells (fig. 35a) showed a moderately recessed,
rather long barrier that at most extended one-third of
the apertural width.
It is obvious from the above paragraphs that varia-
tions show a mosaic pattern and that the variables are
not correlated in any obvious fashion. Widely umbili-
cated Celebes shells have a strong parietal, whereas
the widely umbilicated New Hebridean specimens lack
all trace of a parietal, and the Lau Archipelago exam-
ples have a parietal in the juvenile stage but not the
adult. Viti Levu examples are narrowly umbilicated
and without a parietal barrier.
Material from the island groups covered in this
monograph were relatively distinct from each other,
but when specimens from extralimital areas were con-
sidered, the differences were duplicated. Only one
example from Borabora (BPBM 152396) was seen. Al-
though broken and with the sculpture worn, the large
parietal, relatively wide umbilicus and rather low
spire are very different from the Rurutu shells. These
have the spire markedly elevated and a correlated
rather tight coiling pattern. In some (BPBM 148559)
the body whorl is abruptly deflected, which greatly in-
creases the H/D ratio and narrows the umbilicus. All
examples had a parietal visible from the aperture.
Samoan specimens have a raised spire, slightly in-
creased descension of the body whorl, a relatively nar-
row umbilicus, fairly high H/D ratio, no visible
parietal, and, in the few translucent shells, no parietal
visible through the shell. A subjective impression is
that they have a slightly tighter coiling pattern than
do the Fiji specimens.
Lau Archipelago shells, particularly the largest
sample (BPBM 167137) from Yangasa Levu, are very
widely umbilicated, large shells, with a slight increase
SYSTEMATIC REVIEW
81
FIG. 37. Australian Discocharopa aperta (MbllendorfD. Station WA-113, Glen Helen opening to Finke Gorge, MacDonnell Range, west of
Alice Springs, Northern Territory. A. Solem and L. Price! 111-12-1974. FMNH 182104: a, top of shell (50x); b, side of shell (54x ); c, base of shell
(49x); d, sculpture on body whorl (505x).
in whorl count and distinctly more crowded radial ribs
on the last portion of the body whorl. Visually they are
very similar to the New Hebridean shells. All juveniles
showed a large parietal barrier that was readily visible
from the aperture. No adults showed any trace of a
parietal. The few Viti Levu shells (BPBM 87930,
BPBM 178489) are much more narrowly umbilicated
and are shaped like the Samoan shells. They lack any
trace of a parietal, although at least one is obviously
subadult.
Specimens from Rurutu, Samoa, and the Lau Ar-
chipelago do have a characteristic appearance and if
mixed in a tray could be separated with only slight
difficulty. When compared with various Indonesian
and Bismarck sets, except for the few Rurutu shells
with extreme body whorl deflection, duplicates can be
found for each of the Pacific Island types. Nomen-
clatural recognition would require designation of con-
vergent, isolated geographic races based on use of one
or two characters and ignoring those that disagreed, or
recognizing probably polyphyletic subspecies with in-
termingled mosaic distributions. Neither alternative
seems really adequate, particularly since only material
from one population has been dissected. Eventual rec-
ognition of races or species may be possible, but not
until anatomical studies have been made and more
data are available concerning the meaning of variation
in the parietal barrier.
Genus Sinployea, new genus
Small to very large Charopidae with SV-j-S'/i (more in plano-
spira) normally to rather tightly coiled whorls. Apex and spire rarely
flat, slightly depressed (lamellicosta, tenuicostata) or barely protrud-
ing (descendens), normally moderately protruded, sometimes ele-
vated more than !4 body whorl width (harueyensix, vicaria, angularis,
recursa). Body whorl descending moderately to drastically (ir-
regularis). Apical sculpture normally of 10-12 prominent to fine spi-
ral cords, rarely averaging less than 9 (tahitiensis, allecta) or more
than 13 (angularis, recursa, irregularis), sometimes (peasei, av-
anaensis) greatly reduced in prominence. Postnuclear sculpture of
extremely fine and crowded (avanaensis) to broader and quite widely
spaced (angularis, tenuicostata), usually protractively sinuated ra-
82
SOLEM: ENDODONTOID LAND SNAILS
dial ribs, rarely (recursa) lost on lower spire and body whorl, some-
times becoming too fine and crowded (harveyensis, irregularis) or
irregular (rudis) to count. Microsculpture of fine radial and finer
spiral riblets, secondary spiral cording present in % of species. Um-
bilicus sometimes widely open (descendens, planospira), rarely
(clinta, clausa) closed or barely perforate, normally V-shaped or U-
shaped, with last whorl decoiling more rapidly, margins rounded.
Shell color flammulated, light horn or reddish brown. Sutures deep,
channeled only in proximo, whorls evenly rounded to strongly com-
pressed laterally above periphery, rarely (canalis) with strong lat-
eral compression, periphery rounded, angulated only in angularis.
Aperture circular to compressedly ovate, inclined 5-35 from shell
axis. Body color white or with gray on head and ommatophores.
Pallial cavity with lobes of kidney subequal in length, generally a
strip of lung roof visible between arms of ureter. Genitalia same type
as in Champa. Penis length 0.30-0.53 shell diameter, proportion-
ately larger only in tahitiensis and montana, internally with vergic
papilla, muscular collar just below verge tip, and large stimulatory
pad near middle portion. Relative size of the penial structures vari-
able.
Type species. Sinployea peasei, new species.
Of the other Polynesian genera with typical
stimulatory pad, the Viti Levu Ba has altered shell
form (fig. 74a-c), pallial structures (fig. 75a), and is an
experiment in whorl count reduction. The Tongan
Tuimalila is much larger (table XXXIX), has looser
whorl coiling (fig. 78b, e), altered sculpture, and is
much larger. Maafu (fig. 76a-c) has not been dissected,
but its grossly altered shell sculpture and form im-
mediately distinguish it from any Sinployea. All of the
above groups appear to be specialized derivatives from
Sinployea. In contrast, the genera with apertural bar-
riers, Lauopa, Vatusila, Lagiuala, and Graeffedon,
probably are much more distantly related. Graeffedon
(fig. 87c) has a very distinctive epiphallic structure.
Only a partial specimen of Vatusila tongensis could be
dissected (fig. 84a-b). It seems closely related to
Sinployea.
Intraspecific variation is large enough and
similarities between species sufficiently numerous
that preparation of a key to all species would have
required multiple entries. The more practical solution
of preparing keys to taxa from limited segments of the
generic range has been adopted. Discussions and either
geographic or formal dichotomous keys have been pre-
pared for the species from Society; Cook; Samoan clus-
ter plus Swains Island; Tonga, Ellice, Rotuma, Kusaie
in the Carolines, and Saipan in the Marianas; Fiji; and
Melanesia (New Hebrides, Solomon Islands, Bismarck
Archipelago). In the individual species discussions,
cross-references are made to morphologically similar
forms from other areas, but the main emphasis is on
differentiating taxa from their sympatric or neighbor-
ing species.
Some general discussion of variation patterns is
presented here, although Sinployea is a relatively
uniform group in terms of structure.
Sinployea has the widest geographic range and
greatest number of species recorded for any Polynesian
endodontoid genus. I discuss 49 named species-level
units below and mention two additional taxa rep-
resented by material that is inadequate for description.
Unquestionably, additional species remain to be dis-
covered. Further collections in Tonga, Fiji, Melanesia,
and probably on Tahiti should be especially productive.
Dissection has confirmed placement of the New Hebri-
dean S. euryomphala and the Caroline Island S.
kusaieana in this genus, but inclusion of the Solomon
Island and Bismarck Archipelago species might not be
correct. None of them have been dissected. A record
from Saipan in the Marianas is questionable. It is pos-
sible that Sinployea occurs in New Guinea or even
Wallacea, since Pilsbrycharopa kobelti (Bottger) and P.
brunnescens (Mollendorff) might be classified in
Sinployea. Without dissection, their position will re-
main uncertain. Similarly, some of the New Caledo-
nian species listed in Andrefrancia by Solem (1961, pp.
454-463) possibly might be Sinployea, although I con-
sider this unlikely, since several dissected Andrefran-
cia have distinctive anatomy. Determination of the
exact western distributional limits is beyond the scope
of this study and would require collection of additional
preserved material.
The great Polynesian distribution of Sinployea
probably can be explained on the basis of ecology.
Many species are semiarboreal in habitat, ascending
trees to at least 15 ft., and specimens have been found
frequently in leaf axils or attached to loose bark on stilt
roots. The opportunity for rare accidental transport
during a severe storm is infinitely greater for a species
on loose bark than for one in the dense ground litter.
Correlated with this habitat difference is the develop-
ment of gray markings on the ommatophores, head,
neck, and sometimes mantle edge in those species that
have been found on trees. No trace of gray coloration is
seen on those species living only in ground litter or
under logs and stones. All of the Society Islands and
Cook Island species collected alive in this century show
the darker markings. The condition in many Raroton-
gan species collected on the ground by Garrett, but not
found since the 1880s, is unknown. Of the Samoan
species for which soft parts are known, only the Swains
Island S. intermedia lacks gray coloration. The atoll
environment with its thinner vegetation cover may
have caused this species to become a litter dweller. In
contrast, the Caroline Island S. kusaieana, Fijian S.
inermis, S. adposita, and S. irregularis plus the New
Hebridean S. euryomphala lack all traces of gray col-
oration and have been recorded only from the ground
stratum. Two Lau Archipelago species in particular
may be semiarboreal, but they are known only from
empty shells (S. recursa, S. angularis).
Presently available data suggest that there are
two main centers of speciation, Lau Archipelago (six
species) and Rarotonga (11 species). In both areas there
is evidence of a radiation encompassing both semiar-
boreal and ground stratum niches. It is quite possible
(p. 97) that the Rarotonga litter species are extinct, but
I suspect that the Lau Archipelago habitats may be
less disturbed. Secondary centers of speciation are
found in Samoa (five species) and Tahiti (six species).
SYSTEMATIC REVIEW
83
Elsewhere, only single species on several islands
(Tonga) or separate species on individual islands (Sol-
omons) have been recorded. On Tahiti, only one species
has been collected at a station, and on Rarotonga,
fragmentary data from recent collecting and records in
Garrett (1881) suggest that most, if not all, of the 11
Sinployea species were geographically isolated. In
Samoa and the Lau Archipelago there is ample evi-
dence of rather extensive sympatry. The above specia-
tion pattern suggests a group of relatively recent arri-
val that is well adapted for rare accidental transport
over great distances, but on particular islands tends to
break up into geographic isolates.
In several cases it was possible to dissect sympat-
ric species. In other situations I could study allopatric
species populations from only a few hundred yards
apart. In each situation there have been shifts in size
and/or relative development of the functioning penial
elements (verge, muscle collar, and stimulatory pad)
that probably serve to enable species recognition and to
maintain species isolation. Details of variation in each
situation are given in the discussion of Tahitian
species (p. 85), Cook Island (p. 95), Samoan (p. 118),
and Fijian (p. 147) taxa. Except for these situations of
sympatry or almost certain contact between species, I
found only slight anatomical differentiation. Varia-
tions in whorl count and body whorl capacity affect the
relative proportions of the kidney and whether or not a
strip of lung roof is visible between the ureter arms.
Neither available time nor available material was suf-
ficient to make detailed comparisons of genital propor-
tions. No obvious differences were noted. Although
penis length ranged from 0.8-2.2 mm., when allowance
is made for differences in adult size by calculating the
ratio of mean penis length/mean shell diameter of dis-
sected specimens (table XIV), obviously there is rela-
tively minor variation. Fourteen of 21 dissected
Sinployea have a PL/XD ratio between 0.35 and 0.47. 1
consider it significant that departures from this pat-
tern are shown on the opposite ends of distribution.
The New Hebridean S. euryomphala (Solem), Fijian S.
irregularis (Garrett), and Society Island S. neglecta
have small penes, with a ratio of 0.30-0.31. In contrast,
two Tahitian species found in close proximity have
proportionately very large penes, with ratios of 0.61
and 0.66. Selection pressure for change would occur
only where species came into contact or were sympat-
ric, so that the stable genital pattern is not unexpected.
The apparent correlation between white body color-
litter dweller, gray head and ommatophores-tree trunk
habitat was mentioned above.
Conchological patterns of variation are somewhat
more extensive, but appear mosaic in nature rather
than geographic. Although the range of variation in
most measured characters is rather large, the average
pattern is relatively conservative. Table XV sum-
marizes mean measurements, giving the minimum, 1st
quartile, median, 3rd quartile, and maximum means
for 50 taxa (all described material and the Borabora
TABLE XIV. - RATIO OF PENIS LENGTH TO SHELL DIAMETER IN
SINPLOYEA.
Species
Penis length
in mm.
X Shell
Diameter
PL/D
tahitiensis
2.
1.
2-2.
4-1.
4
5
3.50
3.25
0.66
0.45
lamellicosta
modicella
1.
1.
0-1.
4-1.
1
6
3.35
2.83
0.31
0.53
neglecta
montana
1.
4-1.
65
2.50
0.61
atiensis
1.
1.
1-1.
1-1.
3
3
2.70
2.95
0.44
0.41
andrewi
peasei
1.
1-1.
3
3.15
0.38
avanaensis
1.
4-1.
1.2
6
3.21
2.53
0.47
0.47
clista
aunuuana
1.0
2.76
0.36
allecta allecta
1.3-1.
1.2-1.
1.9-2.
5
6
2
3.13
3.49
4.92
0.45
0.40
0.42
intermedia
complementaria
vicaria vicaria
1.3
1.0
2.86
2.60
0.45
0.38
kusaieana
inermis inermis
1
,8-1.
,8-1.
.65
2.60
2.36
3.29
0.35
0.38
0.50
i . lakembana
adposita
irregularis
.8-1.
0.8
2.87
2.64
0.31
0.30
euryomphala
adult). For the primary measurements of height,
diameter, whorls, and rib count, the 2nd and 3rd quar-
tiles cover from 12%-30% of the total range. Only a few
species are strikingly distinctive in size or shape, most
are very similar.
Four of the seven smallest species were from the
Solomons and Bismarcks, S. nissani, S. kuntzi, S. de-
scendens, and S. novopommerana. The other three
were S. ellicensis nukulaelaeana from Nukulaelae, El-
TABLE XV. - MEAN SPECIES MEASUREMENTS IN SINPLOYEA.
Shell Height In mm.
Shell Diameter in mm
H/D f.atio
Whorl Count
Diameter /Umbilical
Width Ratio
Ribs on Body Whorl 1
MINIMUM
1.00
2.08
0.463
3 1/2
2.73
38.0
1ST QUARTILE MEDIAN 3RD QUARTILE MAXIMUM
Ribs/mm, on Body Whorl 1 3.35
Apical Cords 8.60
1.38
2.50
0.505
3 7/8+
3.69
77.1
8.04
9.40
1.55
2.84
0.538
4 1/8-
4.29
92.5
10.8
10.2
1.77
3.35
0.573
4 3/8-
4.99
112.2
12.2
11.8
2.55
4.92
0.625
6 5/8
CLOSED
225.8
21.0
18.5
1. Four species have reduced ribbing and were omitted
from this table.
84
SOLEM: ENDODONTOID LAND SNAILS
lice Islands, S. godeffroyana from Viti Levu, and the
very modified S. angularis from Namuka, Lau Ar-
chipelago. The largest species is the Upolu Island S.
complementaria, with nine of the 11 Rarotongan
species comprising the other species averaging more
than 3.60 mm. in mean diameter. Concentration of the
smaller species in Melanesia may indicate competition
from terrestrial helicarionids in the 2.5-3.5 mm. size
range, whereas the development of larger Sinployea on
Rarotonga may be the result of that island lacking ter-
restrial helicarionids. Because the Rarotongan radia-
tion of Sinployea probably is extinct, fieldwork to check
these possibilities may be 50 years too late.
Height is closely correlated with diameter in terms
of large and small species, although H/D ratio diverges
in special circumstances. The great increase in whorl
count found in S. planospira (fig. 46d) has resulted in a
lowered H/D ratio, whereas the flat spire, wide um-
bilicus, and slightly increased whorl count in S. canalis
(fig. 49a-c) combine to reduce the H/D ratio. In con-
trast, the nearly closed umbilicus, strongly elevated
spire, and thicker body whorl of the Samoan S. clausa
(fig. 51d-f) give it the highest H/D ratio in the genus.
Only the Tahitian S. montana, with a slightly more
open umbilicus, equally protruded spire, but more
sharply deflected body whorl, is similar in H/D ratio
(fig. 41a-c). The Viti Levu S. irregularis has a flat or
only slightly elevated spire, but a high H/D ratio is
produced solely by an often drastic deflection of most or
all of the body whorl (fig. 65d-f).
Whorl count is a much less variable character. Ig-
noring S. nissani and S. montana, for which only sub-
adult or barely adult material was available, and S.
planospira (fig. 46d), which is characterized by an ab-
normally high whorl count, mean whorls range from
3%- to 5-. Of these, only five of the very largest
species average more than 4 l /2 whorls. There are no
species I can cite as being experiments in whorl count
reduction, whereas only S. planospira shows clear evi-
dence of whorl increase. Its mean whorl count of 6%
includes a juvenile shell and is thus quite conservative,
yet this species averages 1% whorls more than any
other Sinployea.
Relative umbilical width, as measured by the D/U
ratio, is determined by the pattern of coiling, number
of whorls, body whorl contour, and body whorl descen-
sion. Those with wide, cup-shaped umbilici are found
in the Bismarcks (S. novopommerana, fig. 73c),
Rarotonga (S. canalis, fig. 49c; S. planospira, fig. 46f),
Ellice Islands (S. pseudovicaria, fig. 61c), whereas the
very narrowly umbilicated forms are concentrated in
Samoa (all the small species) and Lau (S. angularis,
fig. 64f; S. recursa, fig. 64c; S. adposita, fig. 69f). I do
not know the reason for this.
Whorl contour is relatively stable, with the chan-
neled suture of S. proxima (fig. 46a), strong lateral
compression in S. canalis (fig. 49b), weak supra-
peripheral sulcus in S. rudis (fig. 47e) and S. ir-
regularis (fig. 65e), and angulated periphery in S. an-
gularis (fig. 64e) providing the only major alterations.
Most species vary in the narrow range from an evenly
rounded periphery, as in S. ellicensis (fig. 62b) and S.
vicaria (fig. 58b), to taxa with rather prominent lateral
compression above the rounded periphery, such as S.
recursa (fig. 64b). When specimens are compared di-
rectly, such minor differences are obvious, but no
quantification of these changes was attempted.
Shell ornamentation is relatively complex and the
variations numerous. In several species no counts of
major radial ribs were possible. The Rarotongan S.
rudis (fig. 47d) and the Lau Archipelago S. recursa (fig.
64a) have widely spaced major radial ribs on the upper
spire that become reduced to absent by the body whorl.
In contrast, the Rarotongan S. harveyensis (fig. 48a-b)
and Viti Levu S. irregularis (fig. 65d-e) have the ma-
jor ribs become so fine and crowded on the body whorl
that differentiation of ribs and growth irregularities
becomes impossible. Specimens with few radial ribs in-
clude the very small S. angularis (fig. 64e) and S. in-
ermis lakembana (fig. 66e), the very large S. tenuicos-
tata (fig. 50d), and the larger than average S. adposita
(fig. 69e) and S. lamellicosta (fig. 40a). By far the
greatest number of ribs, mean 225.8, is found in S.
auanaensis (fig. 45e) from Rarotonga. Of the smaller- to
medium-sized species, only the Fijian S. godeffroyana
(fig. 69a) and Lau Archipelago S. navutuensis (fig. 68e)
have actual rib counts equaling some of the larger
Rarotongan shells. The latter three species have by far
the highest ribs/mm, index, whereas S. adposita and S.
tenuicostata have the most widely spaced ribs. The
number of microradial riblets varies with the rib spac-
ing and actual sculpture size. Development or loss of
secondary spiral cording and relative spacing of the
cording showed no consistent pattern that I could re-
late to size or major sculpture.
Apical cording is variable in prominence; the con-
trast between the large cords inS. modicella (fig. la-b)
and the faint ones in S. peasei (fig. 2a-b) indicate the
extent of variation. Actual numbers of cords vary from
a mean of 8.6 in the Samoan S. allecta allecta to a high
of 18.5 in the Viti Levu S. irregularis. There is no sim-
ple correlation between the number of cords and their
relative prominence, because the cord spacing is an
additional variable factor. It was not practical to pre-
pare scanning electron microscope photographs of more
than a few species. The apical cording is too fine for
accurate measurement with a dissecting microscope,
and the relief is too great for effective study with a
compound microscope. Thus, no detailed analysis of
apical sculpture spacing and cord width was possible.
Those species averaging 8.6-9.0 apical cords were S.
allecta allecta from Samoa, S. tahitiensis from Tahiti,
S. decorticata from Rarotonga, S. inermis from Lau,
and S. princei from Taveuni. Those with an enlarged
number of apical cords are clustered geographically.
The Society Island S. montana, S. lamellicosta, and S.
neglecta average 12.5-13.0 cords, whereas the Lau Ar-
chipelago S. adposita, S. inermis lakembana, S. an-
SYSTEMATIC REVIEW
85
gularis, and S. recursa average 13.0-15.9 apical cords.
The presence of both low and high counts in Lau and
Fiji emphasizes the complexity of speciation patterns
in that area.
SOCIETY ISLAND Sinployea
It is rather surprising that no species are known
from Tahaa, Raiatea, or Maupiti. Suitable habitats
were investigated by both Garrett and members of the
Mangarevan Expedition. Only one specimen, too worn
to merit description, has been found on Borabora. Pos-
sibly Sinployea does not live on Maupiti or the Tahaa-
Raiatea complex. Because the Society Islands repre-
sent the eastern limit of distribution for Sinployea,
patchiness of occurrence would not be unexpected.
Single species are found on Huahine and Moorea;
three species, on Tahiti. Because upland collecting on
Tahiti has been limited to two partial transects on Mt.
Aorai and Mt. Orofena, additional taxa may be found
later. Sinployea neglecta (fig. 41d-f) from Huahine
comes the closest of any species in the genus to being
average in characteristics. Its mean measurements
(table XVI) depart from the median Sinployea figures
(table XV) only in having a few more crowded major
radial ribs, more spiral apical cords, and a slightly re-
duced number of microradial riblets. Sinployea mod-
icella (fig. 38a-c) from Moorea is much larger, more
widely umbilicated, and averages fewer apical cords.
There is also a difference in penial size, with S. neg-
lecta having a penis length of 1.4- 1.6 mm. (53% of shell
diameter, which averages 2.83 mm.) and S. modicella
having a penis length of 1.0-1.1 mm. (31.4% of shell
diameter, which averages 3.35 mm.).
All three Tahitian species were collected by the
Mangarevan Expedition during the Mt. Aorai transect
of the malacological assistants. None were taken dur-
ing the Mt. Orofena journey by the expedition bot-
anists and entomologist. Sinployea tahitiensis occurred
in sparse numbers at several stations from 1,400-6,300
ft. elevation and in Fautaua Valley at 250 m. eleva-
tion. Sinployea montana was found at Station 866, a
valley to the west of the Aorai Trail at 6,000 ft. eleva-
tion; S. lamellicosta , at Station 870, a valley west of the
Aorai Trail at 5,000 ft. elevation. Possibly the latter
two were collected in the same valley, but obviously at
different elevations.
Sinployea montana (fig. 41a-c) is a very small,
elevated species with quite narrow umbilicus and
rather widely spaced radial ribs. Sinployea lamellicosta
(fig. 40a-c) has a flat or slightly depressed spire, many
apical spiral cords, a wide umbilicus, and very widely
spaced radial ribs. Sinployea tahitiensis (fig. 38d-f) is
fairly large, has a slightly protruded spire, fewer apical
cords, more whorls, and a thicker body whorl. Penial
differences are marked. Sinployea tahitiensis has a
penis length of 2.2-2.4 mm. (65.8% of shell diameter),
and the penis shows no particular modifications. In S.
montana the penis length of 1.4-1.65 mm. is only 61%
of shell diameter, and the vergic papilla is very large
with the muscular collar reduced in size. Sinployea
lamellicosta has a penis length of 1.4-1.5 mm., or
44.6% of shell diameter, and the stimulatory pad is
greatly enlarged (fig. 39h). Thus S. tahitiensis is
differentiated by penis size, whereas S. lamellicosta
and S. montana have penes of the same size, but in the
former there is enlargement of the vergic papilla and
in the latter, enlargement of the stimulatory pad. If the
latter two species do have a zone of sympatry, probably
the changes in penial structure serve to enforce species
isolation. Further collecting is needed in the moun-
tains of Tahiti in order to establish the ecological re-
TABLE XVI. - RANGE OF VARIATION IN SOCIETY ISLAND SINPLOYEA.
M AMP BUCWWiB ur
SPECIMENS
modicella 399
RIBS RIBS/MM.
112.9(89-145) 10.82(8.59-12.21)
60.7(44-71) 5.86(4.67-8.37)
52.4(43-71) 5.44(4.03-7.91)
(Ferussac)
tahitiensis 23
lamellicosta 19
(Garrett)
Borabora
sp.
3
63.0(57-69)
8.04(7.43-8.65)
172
108.6(71-143)
12.23(8.84-16.87)
1
126
14.14
HEIGHT DIAMETER
1.72(1.35-2.04) 3.35(2.99-3.68)
1.75(1.39-2.09) 3.50(2.98-4.04)
1.58(1.35-1.74) 3.25(2.78-3.55)
1.56(1.52-1.59) 2.50(2.45-2.55)
1.51(1.28-1.81) 2.83(2.57-3.26)
1.55 2.86
H/D RATIO
0.514(0.406-0.598)
0.498(0.428-0.541)
0.487(0.454-0.521)
0.623(0.622-0.623)
0.536(0.441-0.632)
0.540
APICAL
CORDS
SPIRE
ELEVATION
BODY WHORL
WIDTH
WHORLS UMBILICUS D/U RATIO CORDS ELEVATION UTnTH SP/BWW
modic. 4 l/4+(4-4 5/8) 0.96(0.72-1.22) 3.48(2.95-4.28) 9.20(7-13) 0.17(0.07-0.23) 1.01(0.89-1.12) 0.166(0.074-0.226)
tahit. 4 3/8-(4-4 5/8) 0.95(0.66-1.12) 3.68(3.38-4.50) 8.92(6-11) 0.10(0.07-0.13) 1.02(0.95-1.05) 0.104(0.091-0.133)
lamel. 4+(3 3/4-4 3/8) 0.99(0.74-1.22) 3.26(2.89-3.87) 12.50(10-16) FLAT 0.86(0.72-0.99) FLAT OR DEPRESSED
monta. 3 5/8+(3 5/8-3 3/4) 0.36(0.30-0.43) 7.24(5.93-8.55) 13 0.15(0.13-0.16) 0.85(0.84-0.86) 0.226(0.220-0.231)
negle. 4 (3 5/8-4 1/4) 0.65(0.53-0.95) 4.39(3.21-5.00) 12.46(10-15) 0.14(0.10-0.23) 0.78(0.72-0.86) 0.180(0.115-0.275)
Borab. 4 0.69 4.14 WORN 0.23 0.89 0.222
86
SOLEM: ENDODONTOID LAND SNAILS
lationships and geographical ranges of these species.
Despite the size and shape differences, the Society
Island species share a characteristic appearance of
very thin major ribs with periostracal lamellar exten-
sions. Most species from other island groups have dis-
tinctively lower and broader radial ribs. Secondary spi-
ral cording is absent in S. montana, present only on the
spire in S. modicella, and moderately to strongly de-
veloped in the other species. Typical flammulated color
patterning is seen in all but S. montana and S.
tahitiensis, which have monochrome shell color. All
species show a light to dark gray coloration on the head
and neck. Sinployea modicella and S. tahitiensis have
been collected on tree trunks or in axils of low trees,
plus the usual log and under stone habitats. Sinployea
neglecta was found only in the ground stratum, but at
these stations no special investigation was made of ar-
boreal habitats.
Because most species are geographically isolated,
no formal key has been prepared. The following synop-
sis provides quick identification of known taxa:
Moorea: S. modicella (Ferussac, 1840)
Huahine: S. neglecta, new species
Borabora: Sinployea sp. (probably new)
Tahiti: S. montana, new species (diameter
less than 2.75 mm., D/U ratio
more than 5.00, H/D ratio more
than 0.600)
S. tahitiensis, new species (spire ele-
vated, about 9 spiral apical cords,
H/D ratio less than 0.550)
S. lamellicosta (Garrett, 1884) (spire
flat or depressed, about 12.5 spiral
apical cords, H/D ratio less than
0.550)
Discovery of additional species is very probable.
Sinployea modicella (Ferussac, in Deshayes, 1840),
Figures la-e, 38a-c, 39a-b.
Helix modicella Ferussac, 1840, Hist. Nat. Moll. terr. fluv., 1, pi.
86, fig. 3; Deshayes, 1850, Hist. Nat. Moll. terr. fluv., 1, pp.
90-91 "sur les hautes montagnes, Taiti" ( = Tahiti, Society Is-
lands); Pfeiffer, 1853, Monog. helic. viv., 3, p. 92; Pfeiffer, 1859,
Monog. helic. viv., 4, p. 86; Pfeiffer, 1876, Monog. helic. viv., 7, p.
149.
Pitys modicella (Ferussac), Pease, 1871, Proc. Zool. Soc. London,
1871, p. 474.
Patula modicella (Ferussac), Schmeltz, 1874, Cat. Mus. Godeffroy,
5, p. 93 Huahine (?), Society Islands; Garrett, 1884, J. Acad.
Nat. Sci., Philadelphia, 9 (1), pp. 28-29 partly.
Helix (Patula) modicella Ferussac, Tryon, 1887, Man. Conchol., (2)
3, p. 38, pi. 8, figs. 28-30.
Endodonta (Charopa) modicella (Ferussac), Pilsbry, 1893, Man.
Conchol., (2) 9, p. 35.
Diagnosis. Shell larger than average, diameter 2.99-3.68 mm.
(mean 3.35 mm.), with 4-4% normally coiled whorls. Apex and spire
slightly to moderately and evenly elevated, last whorl usually de-
scending more to sharply more rapidly, spire protrusion about '&
body whorl width, H/D ratio 0.406-0.598 (mean 0.514). Apical
sculpture of 7-13 (mean 9.2) prominent spiral cords. Postnuclear
sculpture of narrow, prominent, numerous, protractively sinuated
radial ribs, 89-145 (mean 112.9) on the body whorl, whose interstices
are 2-5 times their width. Ribs/mm. 8.59-12.21 (mean 10.82). Mi-
crosculpture of fine radial riblets, 3-7 between each pair of major
ribs, very fine and crowded spiral riblets, plus weak secondary spiral
cording on spire that fades out on body whorl. Umbilicus widely
open, V-shaped, last whorl decoiling more rapidly, contained 2.95-
4.28 times (mean 3.48) in the diameter, margins rounded. Whorl
contours and apertural features without marked change from aver-
age pattern.
Sinployea modicella (fig. 38a-c) is most similar to
S. neglecta (fig. 41d-f) from Huahine andS. tahitiensis
(fig. 38d-f) from Tahiti. The former is distinctly
smaller, has a narrower umbilicus, more and finer
apical cords, and distinctly more prominent secondary
spiral cording. Sinployea tahitiensis has half as many
radial ribs and many more microradial riblets. No ex-
tralimital species are close in size and shape.
Description of figured specimen. Shell larger than average,
with 4 normally coiled whorls. Apex and spire slightly and evenly
elevated, last whorl descending much more rapidly, H/D ratio 0.480.
Apical whorls 1%, sculpture of 13 narrow spiral ribs. Postnuclear
whorls with narrow, prominent, protractively sinuated radial ribs,
107 on the body whorl, whose interstices are 2-5 times their width.
Microsculpture of fine radial riblets, 3-7 between each pair of major
ribs, crossed by very fine and crowded spiral riblets, plus fine secon-
dary spiral cording equal in size to microradials on spire that become
weak to absent on body whorl. Sutures deep, whorls strongly rounded
above, flattened laterally above periphery and somewhat flattened
on basal margin, with evenly rounded outer margin. Color light red-
dish horn, wide slightly darker flammulations visible above
periphery. Umbilicus widely open, V-shaped, last whorl decoiling
more rapidly, contained 3.63 times in the diameter, margins
rounded. Aperture ovate, strongly flattened laterally above
periphery, somewhat flattened on basal margin, inclined about 25
from shell axis. Height 1.55 mm., diameter 3.22 mm.
Range. Moorea, Society Islands.
Material. Moorea: Opunohu Valley on tree
trunks under bark (15 specimens, BPBM 89709);
Faatoai Valley, 1 1 A miles inland (Stations 886, 893,
896, 900) at 150-300 ft. elevation under dead logs and
leaves or on trunk of hau (335 specimens, BPBM
150331-4, BPBM 150375-9, BPBM 150416-21, BPBM
150473-4); Maramu Valley, l l /z-l% miles inland (Sta-
tions 919, 922) at 50-400 ft. elevation under logs and
dead leaves (30 specimens, BPBM 150710-4, BPBM
150754); Tepatu Valley (Stations 927, 929) at 50-800
ft. elevation (7 specimens, BPBM 150872, BPBM
150895-6); Station 478, above Rotonui, Belvidere Out-
look, Uponohu Bay (1 specimen, FMNH 201054).
"Tahiti" (12 specimens, FMNH 46531, RSM).
Remarks. The taxon Helix modicella has been
used as a catchall for the small, barrier-free endodon-
toids of Polynesia and Fiji. In the absence of precise
measurements and with the availability of only frag-
mentary material, such a usage is understandable. Un-
fortunately, it is difficult to fix the identity of this
name. No type material could be located in Paris at the
time of my study trips. All material collected on Tahiti
in recent years differs from specimens in older collec-
tions that were labeled "Tahiti." These shells agree
more with the populations from lowland Moorea col-
lected by the Mangarevan Expedition. The type figures
have proportions that fall within the range of Moorean
material, since the drawing is 9.4 mm. high, 18.5 mm.
FIG. 38. a-c, Sinployea modicella (Ferussac). "Tahiti," Society Islands. FMNH 155940. d-f, Sinployea tahitiensis, new species. Station 863.
Mt. Aorai, 4,700-5,000 ft., Tahiti, Society Islands. Holotype. BPBM 142072. Scale lines equal 1 mm. (SH).
87
88
SOLEM: ENDODONTOID LAND SNAILS
in diameter, with 4% whorls, H/D ratio 0.509, D/U
ratio 4.02 and about 107 ribs on the body whorl.
Because material from old collections agrees with
the Moorean shells, I have no hesitation in restricting
the type locality to Moorea. The figured specimen is
labeled as coming from "Tahiti" and originating in a
collection formed during the middle 1800s. Possibly it
is part of the type set, but there is no evidence concern-
ing this. In 1981, the Ecole National Superiure des
Mines deposited material from the Deshayes collection
in the Museum National d'Histoire Naturelle, Paris.
Included were two lots of Helix modicella from
"Tahiti." There were 8 and 11 shells glued to strips
of cardboard. Through the courtesy of Dr. Simon Til-
lier, it was possible to examine these specimens.
The smaller lot agrees better in color and sculpture
with the figured shell (fig. 38a-c) than the larger lot,
but may not be conspecific. The latter specimens are
much more brightly colored than the former and have
more widely spaced sculpture. Almost certainly they
are one of the other Society Island species. Without
dismounting and extensive cleaning of each shell, de-
termination of their species identity will not be possi-
ble. I have not chosen a lectotype or neotype. I suspect
that the material from the Deshayes collection was ob-
tained subsequent to illustration of the original
monochromatic specimen. The name Helix modicella
Ferussac, 1840, is applied to the lowland Moorean
populations on the basis of the original illustration.
Variation in local populations is summarized in
Table XVII. All these samples were taken between 50
and 800 ft. elevation. Size and shape are practically
identical, with only umbilical width varying signifi-
cantly. Shells from Maramu Valley (Station 919) have
a distinctly narrower umbilicus than those from
Faatoai Valley (Stations 893, 896), with the small
sample from Tepatu Valley (Station 927) intermediate
in width. Although the difference between shells from
Station 896 and Station 919 is statistically significant,
with 24 degrees of freedom, t = 4.9127, I doubt that
there is any systematic difference.
Populations in Faatoai Valley showed considera-
ble variation in percentage of adults. Station 893, from
150-300 ft. elevation, had 20 of 65 (30.8%) adults; from
Station 896 at 300 ft. elevation, there were only 19 of
182 (10.4%) adults. Station 900, also at 300 ft. eleva-
tion, had 11 of 73 (15.1%) adults. I doubt that sampling
bias is involved, but there are insufficient data con-
cerning the stations to suggest what factors might be
responsible.
Description of soft parts. Foot and tail slightly shorter than
shell diameter. Sole undivided, pedal grooves prominent, uniting
across tail, no caudal horn, middorsal groove weak. Head projecting
in front of foot. Gonopore located slightly above and behind right
rhinophore. Body color yellow-horn, usually with gray suffusion on
head and neck.
Mantle collar (MC) with tapered edge, no glandular extension
onto mantle roof. Pneumostome shielded by a weak right mantle
lobe. Anus (A) and external ureteric pore (KX) in typical position just
inside pneumostome.
Pallial region (fig. 39a) extending about % whorl apically. Lung
roof without granulations. Kidney (K) bilobed, rectal lobe distinctly
longer than pericardial, base tapering and extending apicad under
intestinal loop, upper end medially indented. Length of kidney from
anterior end of rectal lobe to base about 1.51 mm. Ureter (KD) follow-
ing edges of kidney arms, a strip of lung roof visible between arms,
opening next to anus. Heart (H) about half length of pericardial
kidney arm. Principal pulmonary vein (HV) angling toward
pneumostome, not conspicuously branched. Hindgut (HG) starting
just above kidney apex, not tapering.
Ovotestis (fig. 39b) a clump of several finger-like lobes lying
above stomach apex parallel to whorl sides, imbedded in digestive
gland. Hermaphroditic duct (GD) as in S. peasei. Albumen gland
(GG) and talon (GT) typical. Prostate (DG) with very few and large
acini opening into tube lying inside uterine wall. Uterus (UT) bipar-
tite, upper chamber thinner walled and narrower, lower with thick
glandular walls.
Vas deferens (VD) much thicker apically, tapering to slender
tube that remains constant in size until entering epiphallus.
Epiphallus (E) without unusual features. Penial retractor (PR) aris-
ing from diaphragm near apex of pallial cavity, inserting as a U-
shaped fan partly around epiphallus-penis junction. Penis (P) exter-
nally and internally the same as S. peasei, length about 1.00-1.10
mm. Atrium (Y) slender, rather short.
Free oviduct (UV), spermatheca (S), and vagina (V) without un-
usual structures.
(Based on BPBM 150416, 4 individuals, largest 3.49 mm. in
diameter with 4% whorls.)
Sinployea tahitiensis, new species. Figures 38d-f,
39c-e.
Diagnosis. Shell large, diameter 2.98-4.04 mm. (mean 3.50
mm.), with 4-4% normally coiled whorls. Apex and early spire
slightly and evenly elevated, body whorl descending more rapidly,
spire protrusion about '& of body whorl width, H/D ratio 0.428-0.541
(mean 0.498). Apical sculpture of 6-11 (mean 8.92) spiral cords.
Postnuclear whorls with comparatively few, narrow, widely spaced,
protractively sinuated radial ribs, 44-71 (mean 60.7) on the body
whorl, whose interstices are 3-5 times their width. Ribs/mm. 4.67-
8.37 (mean 5.86). Microsculpture of fine radial riblets, 8-12 between
each pair of major ribs, crossed by extremely fine and crowded spiral
riblets with widely spaced secondary spiral cords whose interstices
are about twice those of microradials. Umbilicus broadly open, V-
shaped, last whorl decoiling rapidly, contained 3.38-4.50 times
(mean 3.68) in the diameter, with rounded margins. Sutures chan-
neled, whorls strongly rounded above, not compressed on basal mar-
gin.
The slightly elevated spire, more widely spaced
secondary spiral cording, fewer apical cords and
slightly narrower umbilicus combine to separate
Sinployea tahitiensis (fig. 38d-f) from the relatively
similar S. lamellicosta (fig. 40a-c). Both S. modicella
(fig. 38a-c) andS. neglecta (fig. 41d-f) have many more
radial ribs, whereas S. montana (fig. 41a-c) is much
smaller and has a very narrow umbilicus.
Description. Shell large, with 4V4 loosely coiled whorls. Apex
and first whorl and one-half flat, last whorl descending moderately,
H/D ratio 0.478. Embryonic whorls 1%, with 10 relatively prominent
spiral cords. Remaining whorls with protractively sinuated radial
ribs having high periostracal lamellar extensions, 61 on the body
whorl, whose interstices are 3-5 times their width. Microsculpture of
fine radial riblets, 8-12 between each pair of major ribs, crossed by
extremely fine and crowded spiral riblets. Secondary spiral cording
equal in size to microradials, twice as widely spaced. Umbilicus
broadly open, strongly decoiling on last whorl, V-shaped, contained
3.47 times in the diameter. Color nearly uniform light reddish yellow
with darker periostracal coloring on ribs. Aperture ovate, slightly
flattened laterally above periphery, basal margin evenly rounded,
SYSTEMATIC REVIEW
89
TABLE XVII. - LOCAL VARIATION IN SOCIETY ISLAND SINPLOYEA, I.
modicella
Sta. 919
BPBM 150710
Sta. 900
BPBM 150473
Sta. 893
BPBM 150375
Sta. 896
BPBM 150416
Sta. 927
BPBM 150872
tahitiensis
Sta. 867
BPBM 145702-3
Sta. 862
BPBM 145166
NUMBER OF
SPECIMENS
61.56.52
(55-68)
1.94*0.050
(1.78-2.07)
1.56+0.082
(1.38-1.78)
H/D RATIO
1.
.67+0.
(1.
056
42-1,
,89)
3.2910.
(3.
059
11-3
,51)
.508+0,
(0
,0166
,430-0
,553)
1
.66+0,
(1,
044
39-1
,95)
3.2810,
(3,
044
01-3,
54)
,505+0
(0,
,0087
,456-0,
,551)
1
.75+0
(1.
,028
56-2
.02)
3.3410
(3,
,026
.15-3
.61)
,5230,
(0
,0069
,480-0,
,598)
1
.73+0
(1
,037
.36-2
.05)
3.4010,
(3,
035
,18-3
,71)
,508+0.0085
(0.406-0,
,562)
1
, 720.
(1.
,051
,62-1
.79)
3.39+0
(3,
,090
21-3.51)
,509+0
(0
,0214
,471-0,
,546)
3.74+0.110
(3.42-4.01)
3.2810.069
(3.19-3.49)
0.519+0.0065
(0.500-0.541)
0.47510.0173
(0.428-0.509)
D/U RATIO
APICAL CORDS
modicella
Sta.
919
4 l/8+(4 1/8-4 1/4)
.85+0
(0
,037
,72-0
.97)
3.8910
(3
.106
.54-4
.28)
9.00+0.69
(7-12)
Sta.
900
4 l/4-(4-4
3/8)
.94+0,
(0,
018
,85-1
,05)
3.47+0
3.
.054
16-3.
77)
9.0010.54
(7-12)
Sta.
893
4 1/4-K4-4
1/2)
,9810,
(0
017
,86-1,
,15)
3.4010
(3
.043
.06-3
.78)
9.5810.33
(8-13)
Sta.
896
4 l/4+(4-4
5/8)
1
,02+0,
(0,
021
86-1,
,22)
3.3310
(2
.057
.95-3
.79)
9.17+0.33
(7-12)
Sta.
927
4 l/4-(4 1/8-4 1/4)
.9310,
(0,
061
86-1,
05)
3.6410
(3
.162
.32-3.85)
tahitiensis
Sta. 867 4 1/2(4 3/8-4 5/8) 1.04+0.489
(0.92-1.12)
Sta. 862 4 l/8+(4-4 3/8) 0.9310.041
(0.82-1.02)
3.61+0.061
(3.41-3.75)
3.5510.126
(3.38-3.92)
9.50+0.58
(9-10)
7.5010.65
(6-9)
inclined about 25 from shell axis. Height of holotype 1.74 mm.,
diameter 3.66 mm.
Holotype. Society Islands: Tahiti, Station 863,
Mt. Aorai Trail at 4,700-5,500 ft. elevation. Collected
by Yoshio Kondo and Donald Anderson on September
14, 1934. BPBM 142072.
Range. Interior of Tahiti at 770-6,300 ft. eleva-
tion, Society Islands.
Paratypes. Tahiti: Fautaua Valley (Station 20)
at 250 m. elevation (1 specimen, BPBM 136343); right
fork of Papenoo Valley (Station 951) at 1,400 ft. eleva-
tion (2 specimens, BPBM 144969-70); Aorai Trail (Sta-
tion 863) between 2nd and 3rd camps in leaf axils of
ieie (3 specimens, BPBM 145913); ravine on west side
(Station 862) of ridge crest, Mt. Aorai at 4,600 ft. eleva-
tion (6 specimens, BPBM 145166-7); Aorai Trail (Sta-
tions 865, 867) at 4,700-6,300 ft. elevation (10 speci-
mens, BPBM 145296-7, BPBM 145702-3).
Remarks. The flat spire and more numerous api-
cal cords of S. lamellicosta (fig. 40a-c) are the sys-
tematically important conchological characters sep-
arating that species fromS. tahitiensis (fig. 38d-f). The
former was taken by Garrett at relatively low eleva-
tions and by the Mangarevan Expedition in a valley to
one side of the Mt. Aorai Trail. It probably is geo-
graphically isolated from populations of Sinployea
tahitiensis. Sinployea montana also was collected in a
valley to the west of the Mt. Aorai Trail, but at 1,000 ft.
higher elevation than S. lamellicosta. The Moorean S.
modicella (fig. 38a-c) differs primarily in having twice
as many radial ribs on the body whorl and fewer micro-
radial riblets.
The penis (2.2-2.4 mm. long) is much larger than
in either S. lamellicosta (1.4-1.5 mm. long) orS. mon-
tana (1.4-1.65 mm. long). Internally it shows no
differences from the typical pattern.
K KD
FIG. 39. Anatomy of Tahitian and Moorean Sinployea: a-b, Sinployea modicella (Ferussac). Station 896, Faatoai, Moorea. BPBM 150416. a,
pallial region, b. genitalia; c-e, Sinployea tahitiensis, new species. Station 867, Mt. Aorai, Tahiti. BPBM 145703. c, pallial region, d, genitalia. e,
detail of talon-carrefour region; f, genitalia of Sinployea montana, new species. Station 866, Mt. Aorai, Tahiti. BPBM 145401; g-h, Sinployea
lamellicosta (Garrett). Station 870, Mt. Aorai, Tahiti. BPBM 145537. #, genitalia. h. detail of stimulatory pad in penis. Scale lines equal 1 mm.
(a-d,f-g, MO;e, h, MM).
90
SYSTEMATIC REVIEW
91
Data on material from two local populations are
presented in Table XVII. Specimens from Station 862
were subadult. This lower H/D ratio reflects the les-
sened body whorl descension as does the much lower
mean height. The umbilical width is obviously less
than in specimens of S. lamellicosta in the same size
range. This provides a ready means of separating
specimens where the spire protrusion is not obvious.
Description of soft parts. Pallial region (fig. 39c) unusual only
in having rectal and pericardia! kidney arms equal in length, about
1.38 mm. long. Genitalia (fig. 39d) with head of talon less globular
(fig. 39e) and a distinct shaft. Length of penis stalk, vagina, and
atrium relatively short. Total length of penis about 2.2-2.4 mm.
Interior of penis without unusual features. Jaw with plates partly
fused. Radula typical, 6 laterals, with central slightly smaller.
(Based on BPBM 145703, 3 partial individuals.)
(Garrett, 1884). Figures
Sinployea lamellicosta
39g-h, 40a-c.
Patula lamellicosta Garrett, 1884, J. Acad. Nat. Sci., Philadelphia,
9(1), p. 30, pi. 2, fig. 11 a, b Tahiti, Society Islands.
Helix (Patula) lamellicosta (Garrett), Tryon, 1887, Man. Conchol.,
(2) 3, p. 39, pi. 8, figs. 33-34.
Endodonta (Thaumatodon) lamellicosta (Garrett), Pilsbry, 1893,
Man. Conchol., (2)9, p. 27.
Endodonta (Champa) lamellicostata (sic) (Garrett), Pilsbry, 1893,
Man. Conchol., (2) 9, p. 35 typographical error.
Diagnosis. Shell larger than average, diameter 2.78-3.55 mm.
(mean 3.25 mm.), with 3%-4% relatively loosely coiled whorls. Apex
and spire flat or slightly depressed, body whorl descending sharply,
H/D ratio 0.454-0.521 (mean 0.487), much lower than average. Api-
cal sculpture of 10-16 (mean 12.5) rather fine spiral cords. Postnu-
clear sculpture of prominent, thin, protractively sinuated, widely
spaced radial ribs, 43-71 (mean 52.4) on the body whorl, whose in-
terstices are 4-6 times their width. Ribs/mm. 4.03-7.91 (mean 5.44).
Microsculpture of fine radial riblets, 8-12 between each pair of major
ribs, crossed by extremely fine and crowded spiral riblets that can be
seen on upper spire, plus secondary spiral cording whose spacing is
equal to the microradials. Umbilicus broadly open, wide, cup-shaped,
regularly decoiling, contained 2.89-3.87 times (mean 3.26) in the
diameter, margins rounded. Whorl contours and apertural features
normal.
Sinployea lamellicosta (fig. 40a-c) is the only Soci-
ety Island species with a flat or slightly depressed
spire. Sinployea tahitiensis (fig. 38d-f) is most similar,
but has a slightly elevated spire, narrower umbilicus,
and fewer spiral cords on the apex. All other Society
Island species have much more crowded and numerous
radial ribs. Extralimital species with fiat or depressed
spires all have much more numerous and crowded ra-
dial ribs.
Description. Shell smaller than average, with slightly more
than 3% loosely coiled whorls. Apex and early spire slightly de-
pressed, body whorl descending rather rapidly, H/D ratio 0.440. Em-
bryonic whorls P/K, sculpture partially eroded, but remnants of 11
spiral ribs can be detected. Postnuclear whorls with protractively
sinuated, prominent, widely spaced radial ribs, 50 on the body whorl,
whose interstices are 4-6 times their width. Microsculpture of very
fine radial riblets, 8-12 between each pair of major ribs, crossed by
equal-sized secondary spiral cords that are slightly more widely
spaced than the radial riblets, with barely visible, very fine and
crowded spiral riblets most easily seen on upper spire. Sutures
deeply impressed, whorls strongly rounded above, flattened laterally
above periphery, with evenly rounded basal margin. Color pale
FIG. 40. a-c, Sinployea lamellicosta (Garrett). Tahiti, Society
Islands. Lectotype. BPBM 2841. Scale line equals 1 mm. (MM).
yellow-horn with a few vague, light red flammulations. Umbilicus
broadly open, cup-shaped, regularly decoiling, contained 2.93 times
in the diameter, margins rounded. Aperture ovate, strongly flattened
laterally above periphery, with gently rounded basal margin, in-
clined about 15 from shell axis. Height of lectotype 1.19 mm., diame-
ter 2.70 mm.
92
SOLEM: ENDODONTOID LAND SNAILS
Lectotype. Society Islands: Tahiti. Collected by
Andrew Garrett. BPBM 2841.
Range. Tahiti, Society Islands, at low (?) to high
elevations, probably in only one or two valleys.
Paratype. BPBM 2841.
Material. Tahiti (6 specimens, BPBM 2841, Zu-
rich, NMWC); Pirae-Aorai trail (1 specimen, BPBM
115855 Whitney Expedition); valley west of Aorai
Trail (Station 870) at 5,000 ft. elevation (12 specimens,
BPBM 145537-42).
Remarks. Garrett (1884, p. 30) reported this to
be "a scarce species, living beneath rotten wood in
damp stations." Further collecting is needed to estab-
lish the exact geographic relationships of S. lamel-
licosta, S. tahitiensis, and S. montana in the Mt. Aorai
region.
Whereas the lectotype is the smallest shell ex-
amined and is clearly subadult, the sculptural pres-
ervation was excellent, and it matches in form the
Mangarevan Expedition material. A single juvenile
collected by the Whitney South Sea Expedition (BPBM
115855) is tentatively referred here. The Zurich shells
were barely adult and hence slightly higher and more
narrowly umbilicated than the Mangarevan Expedi-
tion specimens. Only one-third of the latter were adult.
The flat spire and numerous apical cords are the
main features characterizing S. lamellicosta.
Description of soft parts. Pallial region with rectal and pericar-
dial kidney arms almost equal in length, reaching 1.20 mm. long. No
glandular extension onto pallial roof. Genitalia (fig. 39g) with rather
long vagina. Penis quite thick, internal stimulatory pad (fig. 39h)
very prominent. Length of penis about 1.41.5 mm. Glandular zone
of spermatheca with a hardened sperm packet that was 0.45 mm.
long, lower half cylindrical and 0.18 mm. in diameter, upper half
expanded and spherical, diameter 0.28 mm. Radula typical, with 6
laterals.
(Based on BPBM 145537-8, 3 partly extracted specimens.)
Sinployea montana, new species. Figures 39f, 41a-c.
Diagnosis. Shell small, diameter 2.45-2.55 mm. (mean 2.50
mm.), with 3%-3% normally coiled whorls. Apex and spire moder-
ately and evenly elevated, last portion of body whorl starting to
descend more rapidly, spire protrusion between V* and Vs body whorl
width, with a high H/D ratio of 0.622-0.623 (mean 0.623). Apical
sculpture of 13 spiral cords. Postnuclear sculpture of a few thin,
prominent, protractively sinuated radial ribs, 57-69 (mean 63.0) on
the body whorl, whose interstices are usually 3-4 times their width.
Ribs/mm. 7.43-8.65 (mean 8.04). Microsculpture of fine radial rib-
lets, 6-10 between each pair of major ribs, crossed by much finer and
more crowded spiral riblets that become indistinct on body whorl.
Umbilicus quite narrow, U-shaped, last whorl decoiling a little more
rapidly, contained 5.93-8.55 times (mean 7.24) in the diameter,
margins rounded. Basal margin of body whorl rounded, not flattened.
The high spire and very narrow umbilicus of
Sinployea montana (fig. 41a-c) easily separate it from
the other Society Island species. Its rib count is the
same as in S. tahitiensis (fig. 38d-f), but the size
difference produces a much more crowded ribbing in S.
montana. All extralimital species of similar size have
higher whorl counts, lower H/D ratios, and wider
umbilici.
Description. Shell small, with 3% loosely coiled whorls. Apex
and early spire moderately and evenly elevated, last whorl descend-
ing sharply, H/D ratio 0.623. Embryonic whorls 1%, worn traces of
about 13 relatively prominent spiral ribs remaining. Lower whorls
with protractive, thin, radial ribs, 69 on the body whorl, whose in-
terstices are 3-4 times their width. Microsculpture of fine radial
riblets, 6-10 between each pair of major ribs, crossed by much finer
and more crowded spiral riblets that become relatively indistinct on
the body whorl. Sutures deeply impressed, whorls strongly rounded
above, flattened laterally above periphery, evenly rounded on lower
palatal and basal margin. Umbilicus narrowly U-shaped, last whorl
decoiling slightly more rapidly, contained 8.55 times in the diameter,
margins rounded. Color uniform light yellow-horn with darker
periostracal rib extensions, no flammulations. Aperture nearly circu-
lar, slightly flattened laterally above periphery and on basal margin,
inclined about 10 from the shell axis. Height of holotype 1.58 mm.,
diameter 2.54 mm.
Holotype. Society Islands: Tahiti, Station 866,
valley west of Aorai Trail at 6,000 ft. elevation. Col-
lected by Elwood Zimmerman, Yoshio Kondo, and
Donald Anderson on September 15, 1934. BPBM
145401.
Range. Mt. Aorai, 6,000 ft. elevation, Tahiti,
Society Islands.
Paratypes. Tahiti: valley west of Aorai Trail
(Station 866) at 6,000 ft. elevation (2 specimens, BPBM
145401-2).
Remarks. I suspect that the size of Sinployea
montana is underrepresented by the available mate-
rial. Two of the three examples showed only the first
indications of gerontic growth. Probably 10%- 15%
greater diameter is attained.
No other Society or Cook species approaches S.
montana in narrowness of the umbilicus. Only the Sa-
moan S. allecta (fig. 54a-e) and the Fijian S. recursa
(fig. 64c) could be confused on umbilical size. Sinployea
allecta is 0.5 mm. larger and has an average of less
than nine apical cords and a thicker body whorl. Sin-
ployea recursa is still larger and has greatly reduced
major sculpture.
Description of soft parts. Pallial region typical, rectal arm
longer than pericardial, former about 1.25 mm. long in one example.
Apical genitalia not seen, lower portions typical. Penis 1.4-1.65 mm.
long in two examples. Vergic papilla (fig. 39f) large and U-shaped,
muscular bands relatively weak, stimulator typical.
Radula with 5-6 laterals and more than 6 marginals. Central
9 /j. long, 7 ij. wide, distinctly smaller than laterals, 1st lateral 8 n
wide.
(Based on BPBM 145401, 2 extracted and partly broken exam-
ples.)
Sinployea neglecta, new species. Figures 41d-f, 42a.
Diagnosis. Shell of average size, diameter 2.57-3.26 mm.
(mean 2.83 mm.), with 3%-4V4 rather tightly coiled whorls. Apex and
spire slightly to moderately and evenly elevated, body whorl de-
scending more rapidly, spire protrusion Ve- Vs of body whorl width,
H/D ratio 0.441-0.632 (mean 0.536). Apical sculpture of 10-15
(mean 12.5) relatively fine spiral cords. Postnuclear sculpture of fine,
prominent, thin, protractively sinuated radial ribs, 71-143 (mean
108.6) on the body whorl, whose interstices are 2-3 times their
width. Ribs/mm. 8.84-16.87 (mean 12.23). Microsculpture of fine ra-
dial riblets, 3-5 between each pair of major ribs, barely visible mi-
crospiral riblets and relatively prominent secondary spiral cords.
Umbilicus of average width, V-shaped, last whorl usually decoiling
def
f
FIG. 41. a-c, Sinployea montana, new species. Station 866, Aorai Trail, 6,000 ft., Tahiti, Society Islands. Holotype. BPBM 145401: d-f,
Sinplovea neglecta, new species. Station 1008, Tiva, Huahine, Society Islands. Holotype. BPBM 151565. Scale lines equal 1 mm. (a-c, SG; d-f.
MM).
93
94
SOLEM: ENDODONTOID LAND SNAILS
slightly more rapidly, contained 3.21-5.00 times (mean 4.39) in the
diameter, margins rounded. Whorl contours and apertural features
without marked differences from average.
The smaller size, much narrower umbilicus, and
greater number of apical cords separate Sinployea
neglecta (fig. 41d-f) from the Moorean S. modicella (fig.
38a-c). Sinployea sp. from Borabora differs in the
character of the ribbing, which is much lower and more
rounded. Of the Cook Island species, S. andrewi (fig.
44a) has fewer apical cords and more radial ribs,
whereas S. atiensis (fig. 44d-e) is more elevated and
with slightly fewer apical cords. The Tongan S. vicaria
(fig 58a-d) has a U-shaped umbilicus, more rounded
whorl contours, more microradial riblets, and more
whorls.
Description. Shell of average size, with 4 relatively loosely
coiled whorls. Apex and spire moderately and evenly elevated, last
whorl descending slightly more rapidly, H/D ratio 0.530. Embryonic
whorls I'/a, sculpture of about 12 spiral cords. Postnuclear whorls
with numerous protractively sinuated, very thin radial ribs having
periostracal lamellate extensions, 109 on the body whorl, whose in-
terstices are variable, but about 2-3 times their width on middle of
spire. Microsculpture of very fine radial riblets, 3-5 between each
pair of major ribs, with almost equally large, slightly more widely
spaced secondary spiral cords, and barely visible microspiral riblets.
Sutures deeply impressed, whorls strongly rounded above, flattened
laterally with rounded periphery and somewhat flattened basal
margin. Color light reddish horn with indistinct darker reddish
maculations visible mainly above periphery. Umbilicus relatively
narrow, last whorl becoming more strongly decoiled, contained 4.15
times in the diameter. Aperture ovate, flattened laterally above
periphery and on basal margin, inclined about 20 from the shell
axis. Height of holotype 1.45 mm., diameter 2.73 mm.
Holotype. Society Islands: Huahine, Station
1008, 100 yd. inland, near Tiva at 6 ft. elevation. Col-
lected on logs and dead leaves by Yoshio Kondo and
Donald Anderson on October 3, 1934. BPBM 151565.
Range. Huahine, Society Islands.
Paratypes. Huahine (27 specimens, BPBM 6078,
ANSP 1939, ANSP 47759, SMF 165460, ex SMF
165351): Tiva (Station 1008) about 100 yd. inland at 6
ft. elevation on logs and dead leaves (80 specimens,
BPBM 151565-9); Fare (Stations 962, 963) at 100-300
ft. elevation under stones and logs or on dead leaves (52
specimens, BPBM 151121, BPBM 151144-9); Fare
(Station 978) at 1,000-1,200 ft. elevation in a damp
valley on dead fallen leaves (1 specimen, BPBM
151284). "Tahiti" (11 specimens, RSM).
Remarks. Garrett (1884, p. 28) lumped this
species with S. modicella (fig 38a-c). Sinployea ne-
glecta (fig. 41d-f) is much smaller, has more apical
cords, a narrower umbilicus, and more prominent sec-
ondary spiral cording. These differences require better
optical equipment and more precise measurements
than could be made by Garrett. Several small samples
from his collections and two larger sets from the Man-
garevan Expedition are compared in Table XVIII.
There are no significant size or shape differences, ex-
cept in regard to umbilical width. The slightly wider
umbilici for material from Station 1008 and ANSP
47759 is caused by a few gerontic individuals in which
umbilical decoiling has proceeded further than usual,
producing relatively widely open umbilici and thus dis-
torting the means.
Specimens from Tiva (Station 1008) were collected
about 100 yd. inland in coastal vegetation, whereas
those from Fare (Station 963) were found in lowland
forest at 300 ft. elevation further inland. Percentage of
adults was identical, with 15 of 51 (29.4%) from Station
963 and 24 of 81 (29.6%) from Station 1008. Size and
shape differences reached significant levels, particu-
larly in regard to height, with 37 degrees of freedom, t
2.3293, and D/U ratio, t 4.9127. Probably
ecophenotypic variation is involved.
Description of soft parts. Pallial region typical. Rectal kidney
arm 1.45 mm. long, total cavity length 3.3 mm. Narrow strip of lung
roof between ureter arms.
Genitalia (fig. 42a) without unusual features, penis length
1.4-1.6 mm. Drawing misleading in that slender shafts of penis and
vagina folded under. Actual lengths as inS. modicella.
(Based on BPBM 151565, 4 partial examples.)
Sinployea sp.
Material. Society Islands: Borabora, south slope
of Pahio-Temanu ridge (Station 1093) at 800 ft. eleva-
tion (1 specimen, BPBM 152395).
Remarks. A single, probably subadult example
from Borabora is not referable to any named form. The
very low and crowded radial ribbing, 126 ribs on the
body whorl, is more like some Western Polynesian
species than any Society Island Sinployea. Measure-
ments are given in Table XVI. Although obviously
similar to S. neglecta in all dimensions, the character
of the major ribbing is so distinctive that I doubt their
identity. Both have three to six microradials between
each pair of major ribs. Without more material and
knowledge of the apical sculpture, description would be
premature.
COOK ISLAND Sinployea
Specimens of Sinployea have been found on all the
major Cook Islands except Mauke. Libera fratercula
fratercula (Pease) was collected on the latter island by
P. H. Buck, who had taken Sinployea on Atiu,
Aitutaki, and Mangaia during the same expedition.
Hence, there is fair probability that this is an actual
distributional gap.
Sinployea atiensis (Pease) lives on both Atiu and
Aitutaki. Except for a slight increase in whorl count
and a strongly elevated spire (fig. 44e), which increases
the H/D ratio, this is a very average species in all
measured characters (table XIX). Sinployea andrewi
from Mangaia is slightly larger, has a greater number
of more crowded major radial ribs (fig. 44a), is dis-
tinctly less elevated, has fewer microradials, and the
umbilicus is V-shaped, rather than U-shaped.
Rarotonga has, or had, a remarkable radiation of
Sinployea, including nine of the 10 largest species in
the genus. Only the Upolu Island S. complementaria
(Mousson) (table XXIV) reaches larger size than the
Rarotongan species.
SYSTEMATIC REVIEW
95
TABLE XVIII. - LOCAL VARIATION IN SOCIETY ISLAND SINPLOYEA, II.
NUMBER OF
SPECIMENS
H/D RATIO
lamellicosta
Sta. 870
BPBM 145537
50.014.05
(43-57)
1.690.030
(1.62-1.76)
3.49*0.046
(3.38-3.58)
0.485+0.0120
(0.454-0.510)
Sta. 866
BPBM 145401
1.56 0.033
(1.52-1.51)
2.5010.050
(2.45-2.55)
0.62310.0005
(0.622-0.623)
Sta. 1008
BPBM 151565-7
Sta. 963
BPBM 151144, -7
ANSP 1939
ANSP 47759
BPBM 6078
1 .tOZU
(1
,VL.
.29-1
.62)
, /O T U
(2
, uz^
,58-2.
98)
u .
3-41U
(0,
, uuo /
,465-0,
598)
1.5510
(1
,033
,39-1
.82)
2
,8410.028
(2.72-3,
08)
0,
54410.
(0,
0096
500-0.
605)
107.315.36
(101-118)
1.5210
(1
,033
,49-1
.59)
2
.8410
(2
,098
65-2.
98)
0.
53810
(0
.013
.518-0,
563)
107.5112.66
(85-143)
1.5510
(1
.065
,36-1
.69)
2
.9110,
(2
,070
75-3.
08)
0.
54110.
(0,
028
.441-0,
594)
1.6410
(1
.090
,39-1
,99)
2
,9410,
(2.
097
65-3.
28)
0.
55810,
(0.
026
455-0.
632)
lamellicosta
Sta. 870 4 l/4-(4 1/8-4 3/8)
UMBILICUS
1.1410.037
(1.05-1.22)
D/U RATIO
APICAL CORDS
3.0610.0851 11. 25*0. 95
(2.89-3.29) (10-14)
Sta. 866 3 5/8+(3 5/8-3 3/4)
neglecta
Sta. 1008 4-(3 5/8-4 3/8)
Sta. 963 4+(3 7/8-4 1/4)
ANSP 1939 4(3 3/4-4 1/8)
ANSP 47759 4 l/8-(3 7/8-4 1/4)
BPBM 6078 4-K3 7/8-4 1/4)
0.36*0.066
(0.30-0.43)
0.6510.011
(0.59-0.82)
0.63*0.012
(0.56-0.72)
0.6010.011
(0.59-0.63)
0.7210.062
(0.63-0.95)
0.6410.053
(0.53-0.89)
7.24*1.31
(5.93-8.55)
4.2710.051
(3.60-4.72)
4.5310.062
(4.00-4.94)
4.6810.118
(4.45-4.84)
4.0410.222
(3.21-4.53)
4.6410.233
(3.67-5.23)
12.8910.42
(10-15)
12.1310.58
(10-15)
Sinployea peasei (fig. 45a-c) is the Rarotongan
equivalent of S. atiensis and S. andrewi. Reduction of
the apical sculpture (fig. 2a, p. 11) is the significant
change, although (table XIX) there are minor size and
shape differences from the other two species. Sinployea
peasei is replaced in part of Avana Valley by S. av-
anaensis (fig. 45d-f). This is a much more narrowly
umbilicated shell with many very crowded radial ribs
and a lower whorl count. Present data are insufficient
to determine whether there is a zone of sympatry.
There does seem to be ecological separation, because
only one of 159 Sinployea peasei collected in 1964 and
1965 was taken on the leaves of shrubs, the remainder
under logs and stones. A few individuals of S. avanaen-
sis were found under rotting wood, but the great major-
ity were collected on tree trunks in moss and lichens
from 1-15 ft. above ground level. Anatomical dif-
ferences between the two species are slight. Sinployea
avanaensis has the rectal kidney lobe much wider, the
penis about 0.3 mm. longer, and the vergic papilla
longer and narrower than in S. peasei. Study of local
populations in Avana Valley offers an excellent oppor-
tunity for investigation of speciation phenomena.
The larger species show two divergent patterns
that may represent monophyletic series, plus two
species, S. planospira (fig. 46d-h) and S. rudis (fig.
47d-f), that show individual specialization.
One series of species, S. proximo, S. youngi, S.
harveyensis, and S. canalis, retains normal or slightly
more crowded sculpture. The smallest, S. proximo (fig.
46a-c), has a distinctly channeled suture and rela-
tively low spire. Garrett ( 1872, p. 230) reported it was a
"common species lurking under stones and among rot-
96
SOLEM: ENDODONTOID LAND SNAILS
pv
FIG. 42. Anatomy of Society and Cook Island Sinployea: a,
genitalia of Sinployea neglecta, new species. Station 1008, Tiva.
Huahine. BPBM 151565; b, genitalia of Sinployea andreu'i. new-
species. Arakiore Oneroa, Mangaia, Cook Islands. BPBM 97652; c-e,
Sinployea peasei. new species. Station 26, Mt. Maungaroa, 1,400 ft.
elevation. Rarotonga, Cook Islands. FMNH 144645. c, pallial region.
d. penial complex, c, detail of vergic papilla and muscular collar in
penis. Scale lines equal 1 mm. (a-b, MO; e, MM; c-d, PS).
ten wood ... in several villages (error for valleys)."
Later Garrett (1881, p. 386) stated that it was "Not
uncommon in several valleys." A personally annotated
copy of Garrett (1872) corrected "villages" to "valleys."
Sinployea youngi (fig. 48d-f) was taken "On the ground
in damp woods, and only noticed in a single valley"
(Garrett, 1872, p. 229). It is characterized by its large
size, compressed aperture, and widely open umbilicus.
Sinployea harueyensis (fig. 48a-c) is very similar to S.
youngi, but has a higher spire, more numerous crowded
ribs, narrower umbilicus, and ovate aperture. It was "A
common species found under rotten wood" (Garrett,
1872, p. 228). Sinployea canalis (fig. 49a-c) was "found
on the ground in damp forests, and confined to a single
valley" (Garrett, 1872, p. 227). Although S. planospira
(fig. 46d-h) retains typical sculpture, its great increase
in whorl count (mean 6%), very wide umbilicus, flat-
tened apex, plus early spire set it apart from the above
species. Sinployea planospira is "a very rare species"
(Garrett, 1881, p. 388). Its habitat was not defined.
The second series differs primarily in its much
wider spacing of the major ribs. Quite probably it is
polyphyletic. Sinployea decorticata (fig 47a-c) has a
higher spire and H/D ratio with between 10 and 20
microradials between each pair of major ribs and aver-
ages 75 ribs on the body whorl. It is confined to a single
valley (Garrett, 1881, p. 387) where it was very abun-
dant on the ground. Sinployea otareae (fig. 50a-c) has a
lower spire and H/D ratio, averages about 96 ribs on
the body whorl, and has only six to 10 microradial rib-
lets between each pair of major ribs. It lived "under
dead wood on the banks of a stream" in a valley about 3
miles away from that occupied by S. decorticata.
Sinployea tenuicostata (fig. 50d-f) is much larger, has a
flat apex, averages about 51 ribs on the body whorl,
and has eight to 12 rather large microradial riblets
between each pair of majors, indicating an increase in
microradial riblet size and spacing. It was a "somewhat
rare species, having a wide range on the island, and
generally found on the ground on the sides of ravines."
Sinployea rudis (fig. 47d-f) has reasonably typical
upper spire sculpture, but the major ribs become
sharply reduced and irregular on the body whorl,
whereas the lateral flattening above the periphery has
proceeded to form a broad and shallow supraperipheral
sulcus. It is thus an independent pattern of specializa-
tion. Sinployea rudis was a "very abundant species . . .
we gathered hundreds in several valleys" (Garrett,
1881, p. 387). It was taken "on the ground in damp
woods."
Of the large species, four (S. proximo. S. harveyen-
sis, S. tenuicostata, and S. rudis) were widely distrib-
uted or found in several valleys, whereas five (S.
youngi, S. canalis, S. planospira, S. decorticata, and S.
otareae) were found only in a single valley. The niche of
S. planospira was not defined, but all the other species
were stated to occur on the ground or under rotting
wood. Normally Garrett indicated sympatry for
species. Thus, Mautodontha zebrina (Garrett) (see
Solem, 1976b, p. 162) was collected in the same valley
SYSTEMATIC REVIEW
97
FIG. 43. Anatomy ofSinployea avanaensis, new species. Station 32A, Avana Valley. Rarotonga, Cook Islands. FMNH 144671: a, foot length
in relation to shell diameter; b, pallial region; c, genitalia; d, midvisceral genitalia showing uterine sections; e, interior of penis. Scale lines
equal 1 mm. (PS).
with S. otareae. Sinployea youngi was found in the
same valley as S. decorticata, but Garrett did not state
that they were sympatric. He made no other mention of
any Sinployea being collected together. The probability
is that these were geographically isolated.
Use of the past tense is deliberate. Mautodontha
unilamellata (Garrett), Thaumatodon multilamellatus
(Garrett), Libera tumuloides (Garrett), Libera sub-
cavernula (Tryon), Mautodontha zebrina (Garrett), and
the nine large Sinployea were not collected in 1964 or
1965 despite rather extensive coverage of different val-
leys. Overlooking the highly localized species, such as
"one-half acre . . . nearly two miles inland" (Libera
tumuloides), would not be difficult, but the total failure
to find any of the widely distributed species suggests
that actual extinction has occurred. There has been no
recent comprehensive report on the ant fauna of
Rarotonga, but Pheidole megacephala (Fabricius), a
species very destructive to native insects, has been on
Rarotonga since at least 1914 (Wilson & Taylor, 1967b,
p. 47). The same ant is common on Upolu where
Sinployea complementaria (Mousson) still occurs, but
that species is semiarboreal (p. 129) and the shift in
habitat from litter to semiarboreal may have allowed
coexistence of the ant and snail. Possibly some species
still survive at higher elevation in isolated colonies,
but I suspect that most of the Rarotongan endodontoids
are extinct.
98
SOLEM: ENDODONTOID LAND SNAILS
TABLE XIX. - RANGE OF VARIATION IN COOK ISLAND SINPLOYEA.
NUMBER OF
SPECIMENS RIBS RIBS/MM.
atlensls 39 93.3(83-107) 11.27(10.4-12.5)
HEIGHT DIAMETER H/D RATIO
1.57(1.29-1.92) 2.70(2.45-3.18) 0.582(0.527-0.642)
(Pease)
andrewl 83 129.2(104-158) 14.01(11.63-16.09)
1.54(1.29-1.69) 2.95(2.68-3.15) 0.519(0.472-0.568)
pease 1 173 112.2(83-167) 11.51(8.80-14.66)
avanaensis 138 225.8(175-290) 20.82(16.28-25.1)
1.62(1.25-1.94) 3.15(2.97-4.01) 0.492(0.415-0.558)
1.52(1.32-1.81) 3.21(2.86-3.82) 0.474(0.427-0.516)
proxima 20 119.7(92-154) 10.75(8.91-12.96)
(Carrett)
planospira 3 128.0(108-148) 10.62(10.05-11.19)
1.97(1.71-2.37) 3.61(3.26-4.08) 0.544(0.519-0.583)
1.77(1.52-2.22) 3.74(3.44-4.24) 0.468(0.439-0.524)
(Carrett)
rudls 32 RIBS REDUCED
2.23(1.97-2.83) 4.04(3.68-4.83) 0.553(0.500-0.642)
(Garrett)
decorticata 21 75.0(62-103) 5.61(4.84-6.47)
2.36(2.04-2.98) 4.30(3.71-5.10) 0.550(0.488-0.590)
(Garrett )
harveyensis 27 RIBS REDUCED
2.39(1.92-3.01) 4.35(3.91-5.23) 0.550(0.475-0.641)
(Garrett)
young i 17 115.4(96-139) 8.13(7.15-9.68)
(Garrett)
canalls 18 119.8(98-146) 8.22(7.07-9.74)
2.27(1.84-2.66) 4.54(3.75-5.07) 0.499(0.457-0.553)
2.14(1.84-2.63) 4.63(4.01-5.46) 0.463(0.427-0.544)
(Garrett)
otareae 38 95.6(81-116) 6.48(5.72-7.73)
2.41(2.07-2.96) 4.68(4.21-5.20) 0.516(0.470-0.570)
(Garrett)
tenuicostata 15 50.7(43-66) 3.46(2.87-4.23)
2.34(1.94-2.63) 4.70(4.08-5.43) 0.499(0.437-0.537)
(Garrett)
WHORLS UMBILICUS n/U RATIO
atlen. 4 3/8-(4 1/8-4 7/8) 0.63(0.56-0.82) 4.26(3.77-4.82)
APICAL SPIRE BODY WHORL
CORDS ELEVATION WIDTH SP/BWW
11.13(9-14) 0.21(0.16-0.26) 0.86(0.79-0.99) 0.237(0.196-0.296)
andre. 4 l/8+(4-4 3/8) 0.72(0.63-0.79) 4.09(3.74-4.45)
9.19(8-11) 0.17(0.13-0.23) 0.91(0.86-0.95) 0.189(0.143-0.269)
pease. 4 3/8-(4-5 1/8) 0.81(0.56-1.25) 3.93(3.72-4.60)
avana. 4+(3 7/8-4 1/2) 0.69(0.53-0.99) 4.78(3.87-5.69)
11.4(9-14) 0.18(0.10-0.30) 0.93(0.82-1.02) 0.176(0.103-0.300)
10 ~FINE EM 0.17(0.12-0.23) 0.95(0.79-1.09) 0.181(0.113-0.264)
proxi. 4 3/8-(4-4 7/8) 0.68(0.59-0.76) 5.35(4.48-6.16)
piano. 6 5/8(6 1/4-7 1/4) 1.28(1.15-1.41) 2.90(2.75-2.98)
rudls 4 l/4+(3 7/8-4 5/8) 0.83(0.69-1.12) 4.87(4.18-5.62)
0.15(0.10-0.23) 1.17(1.05-1.32) 0.124(0.091-0.175)
0.23 1.02 0.226
10.02(8-12) 0.24(0.20-0.43) 1.32(1.18-1.45) 0.184(0.150-0.296)
decor. 4 5/8-(4 3/8-5) 1.00(0.79-1.22) 4.32(3.75-5.92)
9.00(8-10) 0.27(0.13-0.46) 1.29(1.18-1.58) 0.209(0.103-0.292)
harve. 4 5/8+(4 1/4-5) 0.92(0.76-1.18) 4.73(3.65-5.48)
0.36(0.20-0.59) 1.39(1.28-1.55) 0.258(0.150-0.450)
young. 4 3/8+(3 7/8-5) 1.23(1.03-1.61) 3.73(3.15-4.39)
canal. 5-(4 1/2-5 1/2) 1.59(1.38-1.84) 2.91(2.64-3.10)
10.73(10-12) 0.21(0.15-0.30) 1.39(1.22-1.51) 0.149(0.116-0.196)
10.21(8-12) 0.15(0.10-0.26) 1.53(1.41-1.74) 0.096(0.067-0.151)
otare. 4 3/4(4 1/2-5) 1.15(0.89-1.41) 4.10(3.47-5.52)
9.41(8-12) 0.25(0.13-0.40) 1.45(1.35-1.55) 0.171(0.093-0.261)
tenul. 4 1/4(3 7/8-4 1/2) 0.92(0.79-1.18) 5.16(4.29-6.20)
10.45(9-12) FLAT 1.56(1.44-1.64) FLAT
KEY TO THE RAROTONGAN Sinployea
6. Mean rib count over 200; mid-Avana Valley
1
Whorls less than 5% 2
Sinployea auanaensis, new species
Mean rib count less than 140; widely distributed
?,
Whorls more than 6V 8 Sinployea planospira (Garrett, 1881)
Radial ribs crowded to widely spaced 4
Sinp/oyea peasei, new species
7. Mean ribs/mm, about 8 . . ..8
Mean ribs/mm 37 9
3.
Body whorl with supraperipheral sulcus; ribs reduced to ir-
regularity Sinp!o\ea rudis (Garrett 1872)
8. Umbilicus widely open, mean D/U ratio about 3.00; whorls
strongly compressed laterally
Body whorl without sulcus; ribs very crowded
Sinployea canalis (Garrett, 1872)
4
Sinployea harveyensis (Garrett, 1872)
Suture normal 5
Umbilicus of normal width, mean D/U ratio about 3.75; whorls
Suture distinctly channeled (fig. 46a-b)
Sinployea youngi (Garrett 1872)
Sinployea proxima Garrett, 1872)
9 Spire flat mean rib count about 50
5
Mean diameter less than 3.5 mm 6
Sinployea tenuicostata (Garrett 1872)
Mean diameter more than 4.0 mm.... . ..7
Soire stronslv elevated; mean rib count 75-100.... ...10
SYSTEMATIC REVIEW
99
10. Mean rib count about 75; 10-20 microradials between each pair
of major ribs Sinployea decorticnta (Garrett, 1872)
Mean rib count about 95: 6-10 microradials
Sinployea otareae (Garrett, 1872)
Sinployea atiensis (Pease, 1870). Figure 44d-f.
Pithys atiensis Pease, 1870, J. de Conchyl., 18, pp. 394-395 Atiu,
Cook Islands.
Pitys atiensis Pease, 1871, Proc. Zool. Soc. London, 1871, pp. 453,
474.
Helix atiensis (Pease), PfeifTer, 1876, Monog. helic. viv., 7, p. 165.
Patula atiensis (Pease), Garrett, 1881, J. Acad. Nat. Sci., Philadel-
phia, 8 (4), p. 386 Atiu and Aitutaki, Cook Islands.
Endodonta (Champa) modicella var. atiensis (Pease), Pilsbry,
1893, Man. Conchol., (2) 9, p. 35.
Diagnosis. Shell slightly smaller than average, diameter
2.45-3.18 mm. (mean 2.70 mm.), with 4Vfe-4% rather tightly coiled
whorls. Apex and spire moderately and evenly elevated, slightly
rounded above, last whorl descending more rapidly, spire protrusion
about 'A body whorl width, H/D ratio 0.527-0.642 (mean 0.582). Api-
cal sculpture of 9-14 (mean 11.13) prominent spiral cords. Postnu-
clear whorls with narrow, prominent, crowded, protractively
sinuated radial ribs, 83-107 (mean 93.3) on the body whorl, whose
interstices are 2-3 times their width. Ribs/mm. 10.4-12.5 (mean
11.27). Microsculpture of fine radial riblets, 5-8 between each pair of
major ribs, crossed by very fine and crowded spiral riblets, with a
secondary sculpture of rather widely spaced spiral cords. Umbilicus
rather narrow, U-shaped, regularly decoiling, contained 3.77-4.82
(mean 4.26) in the diameter, margins rounded. Whorl and apertural
characters not differing from average pattern.
Sinployea atiensis (fig. 44d-f) has lower and more
rounded radial ribs than any of the Society Island
species. Its high spire, slightly greater whorl count,
and finer apical cords separate it from S. neglecta (fig.
41d-f). The related S. andrewi (fig. 44a-c) and S.
peasei (fig. 45a-c) are distinctly less elevated and
larger; S. andrewi has fewer and finer apical cords; S.
peasei has the apical cords greatly reduced in promi-
nence.
Types. No specimens dating from Pease's ex-
changes could be located. The possibility of specimens
being discovered in some European museum is good,
and I prefer not to designate a neotype.
Range. Atiu and Aitutaki, Cook Islands.
Material. Aitutaki (5 specimens, Zurich): Reu-
KHI. ] '-2 mile inland at 100 ft. elevation (1 specimen,
BPHM 95649). Atiu (13 specimens, BPBM 87407-9,
Zurich): Matai, Vi mile inland at 50 ft. elevation (11
specimens, BPBM 95117); below Tengatangi, I'/a miles
inland at 150 ft. elevation (1 specimen, BPBM 94942);
Lake Teroto, Vz mile inland at 50 ft. elevation (4 speci-
mens, BPBM 95036); Mokoero, V 2 mile inland at 30 ft.
elevation (2 specimens, BPBM 95141); Tarapaku track,
Vi to '/2 mile inland at 50 ft. elevation (2 specimens,
BPBM 94998).
Remarks. Although Garrett (1884, p. 28) consid-
ered this to be a synonym of the Moorean S. modicella
(fig. 38a-c), the qualitative difference in ribbing and
obvious changes in proportions (tables XVI, XIX) effec-
tively separate the two species. As with the Ancey col-
lection, Pease's types and exchanges were so widely
scattered that I prefer not to select a type specimen at
this time. The Aitutaki examples were barely adult,
and hence I attach no systematic significance to the
smaller size and lower spire shown by this set (table
XX).
Description of. wft parts. Partial dissection of several fragmen-
tary examples showed no differences in structures from S. peasei.
Penis about 1.1-1.3 mm. long, most examples flattened too badly for
opening. One specimen with verge and stimulatory pad identical to
S. peasei.
(Based on BPBM 95117, several examples.)
Sinployea andrewi, new species. Figures 42b, 44a-c.
Diagnosis. Shell slightly larger than average, diameter 2.68-
3.15 mm. (mean 2.95 mm.), with 4-4% normally coiled whorls. Apex
and spire slightly to moderately and evenly elevated, body whorl
descending more rapidly, spire protrusion less than '/s body whorl
width, H/D ratio 0.472-0.568 (mean 0.519). Apical sculpture of 8-11
(mean 9.19) prominent spiral cords. Postnuclear sculpture of narrow,
rounded, crowded, protractively sinuated radial ribs, 104-158 (mean
129.2) on the body whorl, whose interstices usually are less than
twice their width. Ribs/mm. 11.63-16.09 (mean 14.01). Micro-
sculpture of fine radial riblets, 3-6 between each pair of major ribs,
with barely visible microspirals and rather crowded secondary spiral
cording. Umbilicus open. V-shaped, last whorl decoiling more
rapidly, contained 3.72-4.60 times (mean 4.09) in the diameter,
margins rounded. Whorl contour and apertural features normal.
Sinployea andrewi (fig. 44a-c) has a lower spire,
more numerous and crowded radial ribs, and fewer api-
cal cords than S. atiensis (fig. 44d-f). Sinployea peasei
(fig. 45a-c) is larger, with the apical cording reduced in
prominence. Society Island species have much finer
and higher radial ribs. Sinployea rotumana (Smith)
(Fig. 61d-f) has fewer, slightly more widely spaced and
distinctly narrower radial ribs, a U-shaped umbilicus,
and more crowded secondary spiral cording.
Description. Shell of average size, with 4'/4 moderately loosely
coiled whorls. Apex and spire moderately and almost evenly ele-
vated, last whorl descending more rapidly, H/D ratio 0.516. Apical
whorls slightly more than 1%, partly worn, sculpture of 10 spiral
ribs. Postnuclear whorls with closely set, rounded, protractively
sinuated radial ribs, 158 on the body whorl, whose interstices are less
than twice their width. Microsculpture of 3-6 fine radial riblets be-
tween each pair of major ribs, closely set and equally prominent
secondary spiral cords, and barely visible spiral riblets. Sutures
moderately impressed, whorls shouldered above, flattened laterally
above rounded periphery, with slightly flattened basal margin. Color
dark reddish yellow, with occasional lighter streaks. Umbilicus mod-
erately open, V-shaped, last whorl decoiling more sharply, contained
4.13 times in the diameter. Aperture subcircular, strongly flattened
laterally above periphery, slightly flattened basally, inclined about
30 from shell axis. Height of holotype 1.61 mm., diameter 3.12 mm.
Holotype. Cook Islands: Mangaia, Arakiore
Oneroa, % mile from shore at 125 ft. elevation. Col-
lected by P. H. and M. Buck on February 16, 1930.
BPBM 97652.
Range. Mangaia, Cook Islands.
Paratypes. Mangaia: Arakiore Oneroa, % mile
from shore at 125 ft. elevation (82 specimens, BPBM
97652-3); Oneroa, '/2 mile inland under stones at 115 ft.
elevation (1 specimen, BPBM 97544).
Remarks. The appearance of Sinployea andrewi
(fig. 44a-c) is very similar to that of S. neglecta (fig.
41d-f) from Huahine, Society Islands, and it is difficult
to find shell characters that immediately separate in-
dividuals of the two species. In general, S. andrewi has
less sharply impressed sutures and wider, more
crowded radial ribs with a lesser average number of
FIG. 44. a-c, Sinployea andrewi, new species. Arakiore, Mangaia, Cook Islands. Holotype. BPBM 97652; d-f, Sinployea atiensis (Pease).
Matai, Atiu, Cook Islands. BPBM 95117. Scale lines equal 1 mm. (a-c. MM; d-f. SH).
100
atiensis
TABLE XX. - LOCAL VARIATION IN COOK ISLAND SINPLOYEA, I.
NUMBER OF
SPECIMENS
RIBS
HEIGHT
DIAMETER
H/D RATIO
Aitutaki
Zurich
Atiu
BPBM 95117
Zurich
BPBM 87407,
94942, -98,
95036.-141
85.3+1.45
(83-88)
andrewi
BPBM 96652-3
avanaensis
Sta. 18a'
FMNH 144631-2
Sta. 32a
FMNH 144671
24
16
94.42.64
(86-102)
233.3
(175-290)
1.360.038
(1.29-1.42)
1.57+0.034
(1.46-1.79)
1.550.024
(1.49-1.62)
1.640.059
(1.39-1.92)
1.54+0.018
(1.29-1.69)
1.54+0.031
(1.32-1.81)
1.47+0.062
(1.32-1.74)
2.510.039
(2.45-2.58)
2.71+0.063
(2.55-3.18)
2.650.058
(2.50-2.83)
2.750.063
(2.55-3.01)
2.95+0.022
(2.68-3.15)
3.210.060
(2.86-3.68)
3.220.150
(2.89-3.82)
0.5400.0069
(0.527-0.551)
0.5810.0083
(0.543-0.616)
0.5840.&067
(0.573-0.609)
0.596+0.013
(0.539-0.642)
0.519+0.0043
(0.472-0.568)
0.479+0.0062
(0.438-0.516)
0.459+0.0086
(0.427-0.489)
Zurich, AMS
ANSP 47700
BPBM 2317
110.06.60
(92-129)
130.2+8.72
(104-154)
129
(116-142)
1.850.056
(1.72-2.05)
1.96*0.087
(1.79-2.22)
2.090.087
(1.89-2.38)
3.500.068
(3.31-3.68)
3.550.109
(3.28-3.81)
3.79+0.088
(3.61-4.11)
0.5290.0076
(0.518-0.559)
0.5510.0088
(0.534-0.583)
0.5510.0103
(0.523-0.580)
WHORLS
UMBILICUS
D/U RATIO
APICAL CORDS
atiensis
Aitutaki
Zurich 4 l/4-(4 1/8-4 1/4)
0.59+0.010
(0.58-0.61)
4.21+0.012 10.670.335
(4.19-4.23) (10-11)
Atiu
BPBM 4 l/4+(4 1/8-4 5/8)
Zurich 4 l/4+(4 1/8-4 1/2)
BPBM
4 l/2-(4 1/4-4 7/8)
0.640.025
(0.56-0.82)
0.61+0.017
(0.56-0.66)
0.650.019
(0.59-0.72)
4.16+0.090
(3.77-4.59)
4.360.148
(3.87-4.70)
4.35+0.088
(4.05-4.82)
11. 86 0.669
(9-14)
10.50.646
(9-12)
andrewi
BPBM 4 1/8-K4-4 3/8)
avanaensis
Sta. 18a 4+(3 3/4-4 3/8)
Sta. 32a 4 l/8+(4-4 1/2)
proxima
Zurich 4 l/4-(4-4 3/8)
ANSP 4 1/4-K4-4 5/8)
BPBM 4 l/2-(4 1/4-4 7/8)
0.720.010
(0.63-0.79)
0.67+0.025
(0.53-0.86)
0.750.057
(0.63-0.99)
0.650.022
(0.59-0.72)
0.69+0.029
(0.59-0.76)
0.68+0.028
(0.59-0.76)
4.09+0.041
(3.74-4.45)
4.88+0.121
(4.24-5.76)
4.50+0.168
(3.87-4.96)
5.3510.107
(5.00-5.55)
5.160.246
(4.48-5.75)
5.530.206
(4.96-6.16)
9.19+0.203
(8-11)
101
102
SOLEM: ENDODONTOID LAND SNAILS
apical ribs. The differences are sufficient that I can
readily separate individuals on the basis of sculpture,
but I cannot satisfactorily quantify the change.
This species is named after Andrew Garrett,
pioneer collector and student of Cook Island land
shells.
Description of soft parts. Dissection of several individuals
showed no differences from S. peasei. Penis length was about 1.11.3
mm., with internal sculpture typical. Partial figuring (fig. 42b) of the
genitalia is provided.
Radula without unusual features.
(Based on BPBM 97652, 6 individuals).
Sinployea peasei, new species. Figures 2a-b, 42c-e,
45a-c.
Patula atiensis (Pease), Garrett, 1881, J. Acad. Nat. Sci., Philadel-
phia, 8 (4), p. 386 Rarotonga, Cook Islands (partly).
Diagnosis. Shell rather large, diameter 2.97-4.01 mm. (mean
3.15 mm.), with 4-5'/n normally coiled whorls. Apex and spire
slightly and evenly elevated, last whorl descending more rapidly,
spire protrusion a little more than Ve body whorl width, H/D ratio
0.415- 0.558 ( mean 0.492). Apical sculpture of 9- 14 ( mean 11.4) very
faint spiral cords (fig. 2a-b) that cannot be counted on most exam-
ples. Postnuclear whorls with prominent, protractively sinuated,
rounded radial ribs, 83-167 (mean 112.2) on the body whorl, whose
interstices are usually 2-3 times their width. Ribs/mm. 8.80-14.66
(mean 11.51). Microsculpture of fine radial riblets, 48 between each
pair of major ribs, extremely fine spiral riblets, and irregular secon-
dary spiral cording. Umbilicus open, V-shaped, last whorl decoiling
more rapidly, contained 3.72-4.60 times (mean 3.93) in the diameter,
margins rounded. Whorl contours (fig. 2a) and apertural shape typi-
cal.
Sinployea peasei has the apical cording (fig. 2a)
greatly reduced in prominence, but differs only slightly
in size and proportions from the other small Cook Is-
land species. Sinployea modicella (Ferussac) from
Moorea is more widely umbilicated, has finer major
radial ribbing, and fewer, much more prominent apical
cords (fig. Ib).
Description. Shell large, with 4% normally coiled whorls. Apex
and spire slightly and evenly elevated, last whorl descending more
rapidly, H/D ratio 0.495. Apical whorls 1%, sculpture of about 12
spiral cords, partly worn over most of surface. Postnuclear whorls
with prominent, protractively sinuated, lamellate radial ribs, 113 on
the body whorl, whose interstices are 2-4 times their width. Micro-
sculpture of very fine radial riblets, 4-8 between each pair of major
ribs, crossed by extremely fine and crowded spiral riblets, with weak
secondary spiral cording that becomes conspicuous only in the um-
bilicus. Sutures deep, whorls strongly rounded above, markedly flat-
tened laterally above periphery with slightly compressed basal mar-
gin, umbilical margin evenly rounded. Color yellow-horn, with broad
and somewhat irregular reddish flammulations that do not fade out
on shell base. Umbilicus broadly open, V-shaped, last whorl decoiling
more rapidly, contained 3.62 times in the diameter. Aperture ovate,
strongly flattened laterally above periphery, inclined about 30 from
shell axis. Height of holotype 1.71 mm., diameter 3.45 mm.
Holotype. Cook Islands: Rarotonga, Station 26,
west slopes of Mt. Maungaroa at 1,400 ft. elevation.
Collected under dead wood by Laurie Price in De-
cember, 1964. FMNH 155947.
Range. Widely spread over inland areas of
Rarotonga, Cook Islands.
Paratypes. Rarotonga (9 specimens, BPBM 2331,
BPBM 8585): lower slopes of Mt. Maungaroa at 600 ft.
elevation (1 specimen, BPBM 79780); Takuvaine Val-
ley (Station 2) at 200 ft. elevation (1 specimen, FMNH
144519); above Takuvaine stream (Station 4) at 400 ft.
elevation (9 specimens, FMNH 144547); upper
Takuvaine Valley (Station 5a), near Mt. Te Kou at
1,000 ft. elevation (1 specimen, FMNH 144553); Mt. Te
Manga, Takuvaine Valley (Station 30) at 1,200 ft. ele-
vation (12 specimens, FMNH 144644); north slope
(Station 3) of Mt. Maungaroa at 1,000 ft. elevation (9
specimens, FMNH 144536); Mt. Ikurangi (Station 8) at
800 ft. elevation (11 specimens, FMNH 144567-8);
Taipara Valley (Station 10) at 400 ft. elevation (8
specimens, FMNH 144578); west slope of Mt. Maun-
garoa (Station 26) at 1,400 ft. elevation (33 specimens,
FMNH 144645-6); Avana Valley (Station 14) at 300 ft.
elevation (32 specimens, FMNH 144613-4); Avatiu
Valley (Stations 11, 28) at 500-600 ft. elevation (14
specimens, FMNH 144586, FMNH 144652); south
slope of Mt. Te Rua Manga (Station 29) at 1,200 ft.
elevation (9 specimens, FMNH 144654); Rutaki Valley
(Station 13) at 400 ft. elevation (6 specimens, FMNH
144600); Vaikapuangi Valley (Station 24), Mt. Maun-
gatea at 900 ft. elevation (10 specimens, FMNH
144639); summit of Mt. Te Kou (Station R-ll) at 1,930
ft. elevation (3 specimens, FMNH 153388, FMNH
153413). "Cook Islands" (4 specimens, BPBM 167417).
Remarks. Garrett (1881, p. 386) recognized that
the Rarotonga shells were slightly different from the
Atiu and Aitutaki populations (= atiensis), but did not
give them nomenclatural recognition. In naming this
species after W. Harper Pease, I am recognizing his
early encouragement of Andrew Garrett and his de-
scription of many Pacific Island species.
The availability of many localized sets enabled
using this species to establish patterns of variation for
Sinployea and to provide criteria for judging interpopu-
lational variation. As can be seen from Table XXI,
even with only very small numbers of adults from
different stations, there was only minor variability.
Description of soft parts. Foot and tail elongated, slender, not
tapering posteriorly, rounded behind. Sole undivided, without trans-
verse corrugations. Pedal grooves high on foot, uniting across tail,
suprapedal distinctly weaker. No caudal foot or caudal horn. Slime
network rectangular, more prominent on tail than head. Foot trun-
cated anteriorly, head extended well in front. Ommatophores long.
Gonopore slightly behind and above right rhinophore.
Sole and tail yellow-white, head and neck with grayish suffu-
sion. Eyespots black, stalk of ommatophores gray.
Mantle collar (MC) rather short and thick (fig. 42c), with a wide,
even extension of mantle glands onto lung roof. Pneumostome a
single opening masked on each side by swellings of mantle collar.
Anus (A) opening above, but at same level anteriorly as external
ureteric opening (KX). A deep-grooved urinary chamber (LK) leads
into shallow groove from anus through mantle collar.
Pallial chamber (fig. 42c) extending '/2 whorl apically. Lung roof
with heavy concentrations of black speckles along principal pulmo-
nary vein (HV) and mantle glands (MG), with secondary concentra-
tion near hindgut. Kidney (K) U-shaped, rectal lobe distinctly longer
than pericardia!, latter about twice as long as heart (H). Rectal lobe
of kidney lapping completely under hindgut and onto parietal wall.
Base of kidney extended backward between loop of intestine and
outer wall covering, rounded. Ureter (KD) sigmurethrous, U-shaped,
a-f
FIG. 45. a-c, Sinployea peasei, new species. Station 26, Mt. Maungaroa, 1,400 ft., Rarotonga, Cook Islands. Holotype. FMNH 155947; d-f,
Sinployea avanaensis, new species. Station 18A, Upper Avana Valley, 600 ft, Rarotonga, Cook Islands. Holotype. FMNH 155948. Scale line
equals 1 mm. (SH).
103
TABLE XXI. - LOCAL VARIATION IN SINPLOYEA PEASEI.
NAME
NUMBER OF
SPECIMENS
RIBS
RIBS/MM.
HEIGHT
DIAMETER
peasei
FMNH 144547
(Sta. 4)
FMNH 144567
(Sta. 8)
FMNH 144664
(Sta. 30)
FMNH 144536
(Sta. 3)
FMNH 144639
(Sta. 24)
FMNH 144586
(Sta. 11)
FMNH 144652
(Sta. 28)
FMNH 144645
(Sta. 26)
FMNH 144600
(Sta. 13)
FMNH 144654
(Sta. 29)
FMNH 144578
(Sta. 10)
FMNH 144614
(Sta. 14)
BPBM 2331
28
26
113.7+5.17
(93-130)
109.8+7.10
(91-135)
103.7+1.99
(96-116)
108.32.15
(102-115)
106.54.35
(95-115)
106.53.37
(94-119)
124.0+1.33
(100-155)
110. 1+3.07
(83-149)
104.7+7.70
(96-120)
127.215.54
(91-167)
121.3+7.22
(92-156)
114.52.00
(88-141)
11.57+0.478
(9.78-13.24)
11.08+0.366
(10.12-11.90)
11.13+0.186
(10.30-11.89)
10.970.182
(10.39-11.52)
L0.870.413
(9.99-11.59)
10.790.352
9.58-12.12)
11.951.23
(8.80-14.16)
11.55*0.309
(9.13-14.15)
10.49+0.774
(9.39-11.981
12.35+0.842
(10.12-14.42)
11.92+0.473
(10.35-14.66)
11.90+0.209
(9.68-13.51)
1.5010.026
(1.38-1.58)
1.4710.043
(1.38-1.55)
1.3510.022
(1.25-1.45)
1.52+0.058
(1.38-1.84)
1.4610.041
(1.38-1.58)
1.5510.055
(1.38-1.74)
1.7110.132
(1.45-1.84)
1.8410.020
(1.28-1.71)
1.5510.033
(1.51-1.61)
1.62+0.094
(1.41-1.94)
1.6310.054
(1.41-1.84)
1.51+0.020
(1.32-1.71)
1.88+0.070
(1.69-2.12)
3.13+0.045
(3.03-3.36)
3.16+0.148
(2.86-3.85)
2.97+0.056
(2.66-3.16)
3.14+0.058
(2.93-3.32)
3.13+0.112
(2.89-3.42)
3.150.033
(2.99-3.29)
3.340.132
(3.09-3.65)
3.10+0.032
(2.76-3.42)
3.18+0.121
(3.03-3.321
3.24+0.222
(2.80-4.01)
3.22+0.083
(2.83-3.52)
3.0810.030
(2.80-3.45)
3.6110.070
(3.41-3.94)
H/D RATIO
Sta. 4 0.48010.0062
(0.457-0.500)
0.49410.0154
(0.475-0.540)
Sta. 30 0.455+0.0075
(0.415-0.489)
Sta. 3 0.48410.0154
(0.440-0. 554)
Sta. 24 n. 46910. 0051
(0.458-0.478)
Sta.U 0.49110.0145
(0.442-0.546)
Sta. 28 0.51310.0186
(0.468-0.558)
Sta. 26 0.49310.0049
(0.448-0.538)
Sta. 13 0.48710.0235
(0.455-0.533)
Sta. 29 0.50110.0065
(0.484-0.521)
Sta. 10. 0.50610.0092
(0.463-0.535)
Sta. 114 0.49210.0040
(0.447-0.533)
BPBM 2331 0.52110.0166
(0.464-0.587)
WHORLS
4 3/8-(4 1/4-4 1/2)
4 l/4-(4 1/8-4 3/8)
4+(4-4 1/4)
4 l/4+(4 1/8-4 1/2)
4 3/8-(4 1/4-4 1/2)
4 3/8-(4 1/8-4 1/2)
4 5/8+(4 1/4-5 1/8)
4 3/8+(4 1/8-4 3/4)
4 3/8-(4 1/4-4 3/8)
4 l/2-(4 1/8-5 1/8)
4 3/8-(4 1/4-4 1/2)
4 l/4+(4 1/8-4 1/2)
4 5/8+(4 1/2-5)
UMBILICUS
0.7610.021
(0.69-0.86)
0.7910.045
(0.69-0.92)
0.73+0.025
(0.63-0.86)
0.8010.035
(0.72-0.99)
0.7810.043
(0.69-0.86)
0.79+0.015
(0.72-0.86)
0.88+0.068
(0.76-1.05)
0.8110.017
(0.63-0.99)
0.8610.038
(0.79-0.92)
0.8510.109
(0.66-1.25)
0.81+0.039
(0.66-1.02)
0.7910.010
(0.66-0.89)
0.9910.038
(0.82-1.12)
D/U RATIO
4. 1110.059
(3.92-4.38)
4.00+0.092
(3.75-4.18)
4.06+0.081
(3.54-4.29)
3.9410.1211
(3.26-4.21)
4.02+0.1411
(3.69-4.38)
3.97+0.050
(3.80-4.14)
3.83+0.138
(3.47-4.13)
3.88+0.057
(3.33-4.43)
3.7210.090
(3.61-3.83)
3.9210.200
(3.21-4.35)
4.0110.109
(3.45-4.50)
3.90+0.046
(3.52-4.60)
3.6510.144
(3.22-4.36)
104
SYSTEMATIC REVIEW
105
evenly expanded, a strip of lung roof visible between arms of ureter.
At anterior end of rectal kidney lobe, ureter passes alongside hindgut
on lung roof forward to pneumostome. Heart (H) fairly large, angling
slightly from parallel to hindgut. Principal pulmonary vein (HV)
slender, flanked by black speckling, paralleling hindgut until an-
terior quarter of pallial cavity, then angling toward anus, finally
reaching mantle glands. No conspicuous secondary venation.
Ovotestis (G) imbedded in first quarter whorl above stomach-
intestine reflexion, consisting of a single clump of varyingly split
finger-like lobes, internally with iridescent rods. First part of her-
maphroditic duct (GD) a very thin collecting tubule, expanding to a
slender, opaque iridescent tube for length of stomach to middle of
albumen gland (GG), then narrowing to a sinuous narrow tube just
before entering talon (GT). Albumen gland small, darker in color
than prostate (DG), cradled between intestinal loops and base of
kidney in body cavity just above and morphologically below apex of
pallial cavity. Head of spermatheca (S) buried between base of albu-
men gland and head of prostate. Talon a slender tube with bulbously
expanded head down to carrefour, hermaphroditic duct entering lat-
erally on head of carrefour, which tapers from secondary expansion
below apex to head of prostate-uterus. Prostate (DG) short, of large,
white acini entering a groove in uterine wall. Uterus (UT) with 2
chambers, upper (UT,) a very thin-walled sac partly filled with
spongy tissue, opening through a glandular collar into a lower
chamber, white in color, with thick glandular walls, smooth except
for 2 pilasters extending into muscular-walled free oviduct (UV).
Vas deferens (VD) from base of prostate at level of spermathecal
stalk contraction, wide at first with internal pilasters, tapering down
to a thin tube by level of penioviducal angle, passing up along penis,
but not bound to it, until entering epiphallus (E). Latter a swollen
muscular tube with internal longitudinal pilasters entering penis (P)
through a small vergic papilla (PV, fig. 42e). Penial retractor (PR)
arising on diaphragm near apex of pallial cavity, inserting in a U-fan
around base of epiphallus. Penis proper about 1.1-1.3 mm. long,
expanded apically with a minor constriction marking internal mus-
cular collar, then tapering slowly to base of main glandular part,
finally continuing as a slender tube to atrium. Internally (fig. 42d),
penis with a cup-shaped vergic papilla (PV) surrounded by a muscu-
lar collar of 3 circular pilasters. Below this, a huge U-shaped muscu-
lar stimulatory pad occupies rest of penial bulge. Lower part and
section between arms of "U" with walls longitudinally folded.
Atrium (Y) fairly long, without distinctive internal sculpture.
Free oviduct (UV) a short tube with smooth muscular walls ex-
ternally, internally with smooth glandular surface, opening into va-
gina (V) through a glandular collar. Spermatheca (S) and base of
vagina swollen with glandular walls, tapering above to a slender
stalk, then an oval head lying in angle between prostate and albu-
men gland. Vagina (V) long, swollen apically with glandular walls,
lower part a narrow tube with weak pilasters.
Buccal mass and esophagus typical, white salivary glands unit-
ing above esophagus. Stomach expanding just above apex of pallial
cavity, extending % of a whorl apically, occupying parietal and most
of palatal wall surfaces. About V-i of way from apex, stomach narrows
from base, preparatory to stomach-intestine junction. First part of
intestine angling down to columellar margin, running along this
edge to apex of pallial cavity, looping up under kidney to near mid-
point of whorl, reflexing backward for about twice its width, then
turning up to parietal-palatal margin and forward as hindgut. Dis-
tance between apex of pallial cavity and stomach greatly reduced,
loops of intestine very short, hindgut only reaching parietal-palatal
angle at base of kidney.
Free retractor muscle system as in Champa coma.
Enervation of penis from right cerebral ganglion.
(Based on FMNH 144645, several whole examples.)
Sinployea avanaensis, new species. Figures 43a-e,
45d-f.
Diagnosis. Shell rather large, diameter 2.86-3.82 mm. (mean
3.21 mm.), with 3%-4!4 normally coiled whorls. Apex and spire mod-
erately and evenly elevated, last whorl descending more rapidly,
spire protrusion l h to l k of body whorl width, H/D ratio 0.427-0.516
(mean 0.474). Apical sculpture of about 10-14 very fine spiral cords,
rarely distinct enough to be tallied. Postnuclear whorls with very
fine and crowded, protractively sinuated, narrow radial ribs, 175-
290 (mean 225.8) on the body whorl, whose interstices are less than
twice their width. Ribs/mm. 16.28-25.06 (mean 20.82). Micro-
sculpture of fine radial riblets, 2-4 between each pair of major ribs,
barely visible spiral riblets and weak secondary spiral cording visi-
ble on upper surface and basal margins. Umbilicus narrow, V-
shaped, last whorl decoiling more rapidly, contained 3.87-5.69 times
(mean 4.78) in the diameter. Whorl contour typical, aperture a little
more compressedly ovate than normal.
The extremely fine and crowded radial ribs, dark
coloration, and narrower umbilicus effectively sepa-
rate Sinployea avanaensis (fig. 45d-f) from S. peasei
(fig. 45a-c). Other Rarotonga species are much larger
and have fewer or reduced radial ribs. Of the extralimi-
tal species, only the much smaller S. navutuensis (fig.
68d-f) from Lau and S. godeffroyana (fig. 69a-c) from
Viti Levu approach the rib spacing of S. avanaensis. No
other Sinployea has more than 175 ribs on the body
whorl.
Description. Shell rather large, with slightly less than 4'/4
normally coiled whorls. Apex and spire slightly and evenly elevated,
last whorl descending more rapidly, H/D ratio 0.510. Embryonic
whorls 1%, sculpture of about 13 very fine spiral cords, somewhat
worn. Postnuclear whorls with fine, narrow, very crowded, strongly
protractively sinuated radial ribs, about 203 on the body whorl,
whose interstices are no more than twice their width. Microsculpture
of fine radial riblets, 2-4 between each pair of major ribs, crossed by
barely visible spiral riblets, with a weak development of secondary
spiral cording visible on the upper spire, becoming strong on shell
base and in umbilicus. Sutures deep, whorls strongly rounded above,
flatly rounded laterally above periphery, with distinctly flattened
basal margin, umbilical margin strongly rounded. Umbilicus nar-
row, V-shaped, last whorl decoiling much more rapidly, contained
5.33 times in the diameter. Color dark reddish brown. Aperture com-
pressedly ovate, slightly flattened laterally above periphery, dis-
tinctly flattened basally, inclined about 25 from shell axis. Height of
holotype 1.61 mm., diameter 3.16 mm.
Holotype. Cook Islands: Rarotonga, Station 18a,
upper Avana Valley at 600 ft. elevation. Collected by
Laurie Price on lichens and moss-covered branches in
December, 1964. FMNH 155948.
Range. Upper Avana Valley at 300-1,000 ft.
elevation, Rarotonga, Cook Islands.
Paratypes. Rarotonga: Avana Valley between
300 and 1,000 ft. elevation (Stations 18, 18a, 32a, R-13,
R-16) under loose bark or fungus and in crevices on
moss and lichen-covered tree trunks and branches up
to 15 ft. above ground level (137 specimens, FMNH
144631-2, FMNH 144671, FMNH 153381, FMNH
153383-5).
Remarks. At the lowest elevation (Station R-13,
300 ft.) specimens were found only 1-6 ft. above ground
level, whereas at the higher stations examples were
taken up to 15 ft. above the ground. Probably they
move higher, but practical problems in collecting pre-
vented determination of upper limits. All examples
were taken in and under moss and lichens or under
loose bark, except for a few from Station 18 that were
found under dead wood on the ground. Conditions were
quite dry at the time, and the latter occurrence may
represent a partial retreat after moisture. Size and
shape variation is minimal (table XX).
106
SOLEM: ENDODONTOID LAND SNAILS
The very reduced apical sculpture, almost uniform
color, very crowded and numerous radial ribs, much
narrower umbilicus, and more compressedly ovate
aperture immediately separate S. avanaensis from the
common S. peasei. The latter has been taken in lower
Avana Valley (Station 14) at about 300 ft. elevation. It
is widely distributed on other parts of Rarotonga and
has been found on the summit of Mt. Te Kou, which
heads both Takuvaine and Avana Valleys. The two
species are probably allopatric, with S. peasei living
lower in Avana than S. avanaensis.
Description of soft parts. Foot and tail as in S. peasei, length
slightly less than shell diameter (fig. 43a). Sole undivided, pedal
grooves uniting above tail, suprapedal distinctly weaker. No caudal
horn, middorsal groove weak, slime network typical. Gonopore
slightly behind and above right rhinophore. Body color yellow-white,
with gray-black speckling on head and neck region. Mantle collar
with tapered edge, a weak glandular extension onto pallial roof.
Pneumostome shielded by a weak thickening of collar to form a right
mantle lobe. Anus (A) and external ureteric pore (KX) opening at
angle just inside pneumostome.
Pallial region (fig. 43b) extending apically M> whorl, lung roof
clear, without granules. Kidney (K) bilobed, rectal lobe distinctly
longer and broader than pericardial, former lobe crossing hindgut to
reach parietal wall, pericardial lobe about twice as long as heart.
Apex tapering, U-shaped. Ureter (KD) complete, opening alongside
anus, a strip of lung roof visible between ureter arms. Heart (H)
almost parallel to hindgut, proportionately broad. Principal pulmo-
nary vein (KV) angling toward pneumostome, without conspicuous
lateral branches. Hindgut iHG) starting at apex of kidney, following
parietal-palatal margin to anus.
Ovotestis (fig. 43c) occupying slightly less than M> whorl above
stomach reflexion, composed of 5-6 long, single or bifurcated,
finger-like lobes lying parallel to whorl sides, merging irregularly
near base. Hermaphroditic duct (GD) at first a very slender tubule,
becoming abruptly swollen below stomach apex, iridescent until ab-
ruptly narrowing to reflex up albumen gland surface to base of talon.
Albumen gland (GG) oval, deeply indented by head of spermatheca,
intestinal loop, and digestive gland. Talon (GT) with circular head,
hermaphroditic duct entering shaft just below head area, tapering to
head of prostate-uterus. Prostate (fig. 43d) of long, finger-like acini,
sometimes bifurcated, opening into a closed tube contained within
uterine chamber. Uterus (UT) as in S. peasei.
Vas deferens (VD) wide and with glandular walls at first, nar-
rowing before penioviducal angle, becoming a slender tube passing
alongside penis to epiphallus. Latter an enlarged muscular tube with
longitudinal pilasters, ending in a narrow neck before a muscular
collar leading into penis. Penial retractor (PR) arising from dia-
phragm near pallial cavity apex, inserting in a U-shaped fan around
base of epiphallus. Penis (P) about 1.4-1.6 mm. long, swollen on
apical '/2-%, a slight constriction marking end of muscular collar,
lower '/s-'/z a slender tube. Internally (fig. 43e) with a small verge
(PV) nestled in a muscular collar generally containing two circular
bands and followed by the typical doughnut cushion. Lower portion
without distinctive sculpture. Atrium (Y) relatively long, equal in
diameter to basal portion of penis, without special sculpture.
Free oviduct (UV) tapering, internally with glandular tissue and
weak pilasters, opening above end of glandular zone in spermathecal
region. Spermatheca (S) with head lying next to albumen gland,
expanded near base with strong glandular zone that ends abruptly
below junction with free oviduct. Vagina (V) tapering sharply after
end of glandular zone, without special structures present.
Free muscle system as in S. peasei. Buccal retractors not split,
esophagus typical. Stomach extending for % of a whorl, occupying
normal position, expanding and narrowing abruptly. Intestinal loops
compressed into 'Ae of a whorl between pallial cavity apex and
stomach, tapered part of kidney extending well above much of loops.
Digestive glands extending about l*/4 whorls above ovotestis apex,
reduced to narrow strips along stomach, expanding in short area
between stomach and kidney apex.
(Based on FMNH 144671, Station 32a, 4 adults, largest 3.49
mm. in diameter with 4V4+ whorls.)
Sinployea proxima (Garrett, 1872). Figure 46a-c.
Pitys proxima Garrett, 1872, Am. J. Conchol., 7 (4), p. 230, pi. 19.
fig. 24 common in several valleys on Rarotonga, Cook Islands.
Patula proxima (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5,
p. 93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p.
386.
Helix proxima (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p.
543.
Helix (Patula) proxima (Garrett), Tryon, 1887, Man. Conchol., (2)
3, p. 39, pi. 8, figs. 41-43.
Endodonta (Charopa) proxima (Garrett), Pilsbry, 1893, Man. Con-
chol., (2)9, p. 35.
Diagnosis. Shell relatively large, diameter 3.26-4.08 mm.
(mean 3.61 mm.), with 44% normally coiled whorls. Apex and spire
slightly to moderately and evenly elevated, rounded above, last
whorl descending much more rapidly, spire protrusion about 'A body
whorl width, H/D ratio 0.519-0.583 (mean 0.544). Apical sculpture of
12-15 very weak spiral ribs, usually too fine or worn for accurate
counting. Postnuclear whorls with fine, narrow, prominent, strongly
protractively sinuated radial ribs, 92-154 (mean 119.7) on the body
whorl, often irregularly spaced, whose interstices usually are 2-3
times their width. Ribs/mm. 8.91-12.96 (mean 10.75). Micro-
sculpture of fine radial riblets, 3-8 between each pair of major ribs,
crossed by much finer and more crowded spiral riblets. Umbilicus
narrow, U-shaped, last whorl decoiling slightly more rapidly, con-
tained 4.48-6.16 times (mean 5.35) in the diameter, margins
rounded. Whorl contours typical, sutures unique in distinct channel-
ing after first postnuclear.
Sinployea proxima (fig. 46a-c) is immediately rec-
ognizable by its channeled suture. Its reduced apical
sculpture and relatively narrow umbilicus separate it
from most Rarotongan species. Sinployea avanaensis
(fig 45d-f) is similar in umbilical size, but has a normal
suture, much more numerous and crowded radial ribs,
and a distinctly more elevated spire.
Description. Shell larger than average, with 4% moderately
tightly coiled whorls. Apex and spire moderately elevated, rounded
above, body whorl descending distinctly more rapidly, H/D ratio
0.557. Apical whorls 1%, sculpture eroded with traces of weak spiral
ribs remaining. Postnuclear whorls with moderately prominent, very
crowded, protractively sinuated radial ribs, 154 on the body whorl,
whose interstices are about twice their width. Microsculpture of fine,
radial riblets, 2-6 between each pair of major ribs, crossed by
slightly finer and more crowded spiral riblets, with a secondary
sculpture of rather widely spaced spiral cords that become less dis-
tinct on body whorl. Sutures deeply channeled, whorls flatly rounded
laterally above periphery and on basal margin. Color light yellow-
horn with prominent, regular, reddish flammulations. Umbilicus
narrow, U-shaped, slightly and regularly decoiling, contained 5.24
times in the diameter. Aperture ovate, flattened laterally above
periphery and on basal margin, inclined about 20 from shell axis.
Columellar and basal wall with moderate white callus. Height of
lectotype 2.11 mm., diameter 3.79 mm.
Lectotype. Cook Islands: Rarotonga. Collected by
Andrew Garrett. ANSP 47700.
Range. Under stones and among rotting wood in
several valleys on Rarotonga, Cook Islands.
Paratypes.BPBM 2317, ANSP 47700.
Material. Rarotonga (14 specimens, AMS,
Zurich).
a be
def
FIG. 46. a-c, Sinployea proximo (Garrett). Rarotonga, Cook Islands. Paratype. BPBM 2317; d-h, Sinployea planospira (Garrett). g-h,
Details of microsculpture. Rarotonga, Cook Islands. Lectotype. Zurich. Scale lines equal 1 mm. (a-c, SH; d-h, SG).
107
108
SOLEM: ENDODONTOID LAND SNAILS
Remarks. Although Garrett (1872, p. 230; 1881,
p. 386) reported this as a "common" or "not uncommon"
species found in several valleys, no additional material
has been collected. The scattered Museum sets show
clear indication of trading bias (table XX), with the
BPBM residue of Garrett' s personal collection contain-
ing the largest shells, the types in the ANSP only
slightly smaller, and the specimens that eventually
reached Mousson (now Zurich) and Hedley (AMS)
noticeably smaller.
Channeling of the suture is relatively uncommon
in the Pacific Basin endodontoids, being found
elsewhere in the New Caledonian Andrefrancia al-
veolus (Gassies) (see Solem, 1961, pp. 454456, fig. 12)
and the Bismarck Lagiuala macroglyphis (Rensch) (see
p. 188). Ignoring the sutural alteration, S. proximo is
only a slightly enlarged, more narrowly umbilicated
derivative from the peasei stock.
Sinployea planospira (Garrett, 1881). Figure 46d-h.
Patula planospira Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8
(4), p. 388 Rarotonga, Cook Islands.
Helix (Patula) planospira (Garrett), Tryon, 1887, Man. Conchol.,
(2)3, p. 41.
Endodonta (Champa) planospira (Garrett), Pilsbry, 1893, Man.
Conchol., (2)9, p. 35.
Diagnosis. Shell relatively large, diameter 3.444.24 mm.
(mean 3.74 mm.), with 6'/4-7'/4 very tightly coiled whorls. Apex and
early spire flat, body whorl descending rapidly, spire protrusion
about Vs-'/4 body whorl width, H/D ratio 0.439-0.524 (mean 0.468).
Apical sculpture eroded in available material, traces of spiral cord-
ing visible in one umbilicus. Postnuclear whorls with narrow, prom-
inent, protractively sinuated radial ribs. 108-148 (mean 128.0) on
the body whorl, whose interstices are 2-4 times their width.
Ribs/mm. 10.05-11.19 (mean 10.62). Microsculpture of fine radia!
riblets, 5-8 between each pair of major ribs, slightly finer and more
crowded spiral riblets, and secondary spiral cording more clearly
visible in umbilicus. Umbilicus widely open, cup-shaped, regularly
decoiling, contained 2.75-2.98 times (mean 2.90) in the diameter,
margins strongly rounded. Whorls strongly rounded above, flattened
laterally and on basal margin.
Sinployea planospira (fig. 46d-h) has a wide um-
bilicus and much higher whorl count than any other
Polynesian species. Sinployea canalis (fig. 49a-c) is
much larger, has one whorl less, and a much less ele-
vated spire, although it is very similar in rib count and
umbilical proportions (table XIX).
Description. Shell larger than average, with 6'A tightly coiled
whorls. Apex and spire flat, body whorl descending slightly, H/D
ratio 0.442. Embryonic whorls 1%, sculpture mostly eroded, with
traces of widely spaced spiral ribbing remaining. Postnuclear whorls
with narrow, protractively sinuated radial ribs, 108 on the body
whorl, whose interstices are 2-4 times their width. Microsculpture of
very fine radial ribs, 5-8 between each pair of major ribs, crossed by
slightly finer and more crowded spiral riblets with a secondary
sculpture of spiral cording developed in the umbilicus. Sutures deep,
whorls strongly rounded above, flattened laterally and on basal mar-
gin. Umbilicus U-shaped, broadly open, regularly decoiling, con-
tained 2.98 times in the diameter. Color light yellowish white with
traces of irregularly spaced, broad, reddish maculations. Aperture
subtriangular, strongly flattened laterally and on basal margin, in-
clined about 15 from the shell axis. Height of lectotype 1.51 mm.,
diameter 3.42 mm.
Lectotype. Cook Islands: Rarotonga. Collected by
Andrew Garrett. Zoologisches Museum der Universitat
Zurich, Mousson collection ex Museum Godeffroy in
1882.
Range. Rarotonga, Cook Islands.
Paratypes. Rarotonga (2 specimens, Zurich).
Remarks. Because the collections of the Museum
Godeffroy were destroyed during World War II and
Garrett (1881, p. 388) stated that all material of this
species had been deposited in that collection, I had as-
sumed that this was a lost species. The discovery of two
shells in the Mousson collection at Zurich that had
been obtained from the Museum Godeffroy in 1882 al-
lowed recognition of this taxon. Although Mousson had
attached a manuscript name "rarotongensis," I have no
doubt as to their identity. A third shell, also obtained
by Mousson from the Museum Godeffroy, was under
the manuscript name "ortospirata Mousson." It is a
gerontic individual. Unfortunately, the type specimen
was damaged during handling, hence the figures have
dotted lines indicating the upper whorls.
The high whorl count is unmatched by other
Polynesian Charopidae and is much more similar to
the pattern observed in the Endodontidae. Traces of
apical sculpture are sufficiently clear to leave no doubt
as to its affinities. Only S. canalis of the Rarotongan
species has the umbilicus as widely open. Differences
are given in the diagnosis.
Sinployea rudis (Garrett, 1872). Figure 47d-f.
Pitys rudis Garrett, 1872, Am. J. Conchol., 7 (4), pp. 227-228, pi.
19, fig. 18 several valleys on Rarotonga, Cook Islands.
Patula rudis (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5, p.
93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 387.
Patula sublaminata "Mousson" Schmeltz, 1874, Cat. Mus. Godef-
froy, 5, p. 94 Nude name, tentatively synonymized by Garrett
(1881).
Helix rudis (Garrett), Pfeiffer, 1876, Monog. helic, viv., 7, p. 547.
Helix (Patula) rudis (Garrett), Tryon, 1887, Man. Conchol., (2) 3,
pp. 39-40, pi. 8, figs. 44-46.
Endodonta (Charopa) rudis (Garrett), Pilsbry, 1893, Man. Con-
chol., (2) 9, p. 35.
Diagnosis. Shell large, diameter 3.68-4.83 mm. (mean 4.04
mm.), with 378-4% normally coiled whorls. Apex flat, spire moder-
ately and evenly elevated, last whorl descending slightly more
rapidly, spire protrusion more than Vs body whorl width, H/D ratio
0.500-0.642 (mean 0.553). Apical sculpture of 8-12 (mean 10.02)
spiral cords and weak radial corrugations. Postnuclear sculpture of
low, narrow, rather widely spaced protractive radial ribs on upper
spire, becoming reduced and very irregular early on body whorl,
absent near aperture. Microsculpture of fine radial riblets, crossed by
finer and more crowded spiral riblets, reduced on body whorl. Um-
bilicus narrow internally, V-shaped, body whorl decoiling much
more rapidly, contained 4.18-5.62 times (mean 4.87) in the diameter,
margins rounded. Whorls flattened and indented laterally above
periphery, producing a distinct supraperipheral sulcus, strongly
rounded on lower margins, aperture inclined about 35 from shell
axis.
In having a supraperipheral sulcus and losing
major ribbing on the lower spire and body whorl,
Sinployea rudis is quite distinctive among Polynesian
species.
Description. Shell relatively large, with slightly more than 4V4
loosely coiled whorls. Apex and spire slightly but evenly elevated,
body whorl descending more rapidly, H/D ratio 0.531. Apical whorls
l-tL-,
f
FIG. 47. a-c, Sinployea decorticata (Garrett). Rarotonga, Cook Islands. Paratype. BPBM 2313; d-f, Sinployea rudis (Garrett). Rarotonga,
Cook Islands. Paratype. BPBM 2320. Scale line equals 1 mm. (a-c, SH; d-f, MM).
109
110
SOLEM: ENDODONTOID LAND SNAILS
I'/a, sculpture of 11 prominent spiral ribs with a secondary micro-
sculpture of radial and spiral riblets. Postnuclear whorls with nar-
row, low, irregular, protractively sinuated radial ribs, becoming
greatly reduced or absent on the body whorl. Microsculpture of fine
and somewhat irregular radial riblets crossed by distinctly finer and
more crowded spiral riblets that are visible only at 96 x magnifica-
tion under strong lighting. Sutures deep, whorls shouldered above,
flattened to indented laterally above periphery with gently rounded
basal margin. Color greenish yellow-horn with extensive irregular
reddish flammulations. Umbilicus narrow, V-shaped, regularly de-
coiling, aperture contained 5.42 times in the diameter. Height of
lectotype 2.27 mm., diameter 4.28 mm.
Lectotype. Cook Islands: Rarotonga. Collected by
Andrew Garrett. ANSP 47701.
Range. Several valleys on Rarotonga, Cook Is-
lands.
Paratypes. BPBM 2320, ANSP.
Material. Cook Islands (9 specimens, BPBM
167420, FMNH 93820, FMNH 108559): Rarotonga (20
specimens, BPBM 167421, FMNH 46267, ANSP
47701).
Remarks. Garrett (1881, p. 387) indicated that
Sinployea rudis was common and that "we gathered
hundreds in several valleys." In its flat spire, reduced
sculpture, and very prominent supraperipheral sulcus,
S. rudis is immediately separable from the other
Rarotongan species.
The size difference between the type set and
BPBM material (table XXIII) undoubtedly reflects re-
tention of large specimens by Garrett.
Sinployea decorticata (Garrett, 1872). Figure 47a-c.
Pitys decorticata Garrett, 1872, Am. J. Conchol., 7 (4), p. 228, pi.
19, fig. 19 a mountain ravine on Rarotonga, Cook Islands.
Patula decorticata (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy,
5, p. 93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p.
387.
Helix decorticata (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p.
542.
Helix (Patula) decorticata (Garrett), Tryon, 1887, Man. Conchol.,
(2) 3, p. 40, pi. 8, figs. 47-49.
Endodonta (Champa) decorticata (Garrett), Pilsbry, 1893, Man.
Conchol., (2) 9, p. 35.
Diagnosis. Shell very large, diameter 3.71-5.10 mm. (mean
4.30 mm.), with 4%-5 normally coiled whorls. Apex and spire mod-
erately to strongly elevated, body whorl descending more rapidly,
spire protrusion about 1 k body whorl width, H/D ratio 0.488-0.590
(mean 0.550). Apical sculpture of 8-10 (mean 9.0) fairly prominent
spiral cords. Postnuclear whorls with high, thin, prominent, protrac-
tively sinuated and widely spaced radial ribs, 62-103 (mean 75.0) on
the body whorl, whose interstices are 3-4 times their width.
Ribs/mm. 4.84-6.47 (mean 5.61). Microsculpture of very fine radial
riblets, 10-20 between each pair of major ribs, crossed by even finer
and more crowded spiral riblets. No secondary spiral cording. Um-
bilicus narrow, U-shaped, last whorl decoiling more rapidly, con-
tained 3.75-5.92 times (mean 4.32) in the diameter, margins
rounded. Whorl contours and aperture typical.
Sinployea decorticata (fig. 47a-c) is more elevated
and has fewer ribs and double the number of microra-
dial riblets found in S. otareae (fig. 50a-c). Sinployea
tenuicostata (fig. 50d-f) is larger, more depressed, has
fewer ribs, and a flat spire. Other Rarotongan species
differ in their finer and more crowded ribbing.
Description. Shell very large, with 4% normally coiled whorls.
Apex and spire moderately elevated, body whorl descending a little
more rapidly, H/D ratio 0.528. Apical whorls 1%, sculpture of 8 prom-
inent spiral ribs, microsculpture obscured. Postnuclear whorls with
low, rounded, protractively sinuated radial ribs, 73 on the body
whorl, whose interstices are 3-4 times their width. Microsculpture
occasionally visible as fine radial riblets, 10-20 between each pair of
major ribs, crossed by much finer and more crowded spiral riblets.
Sutures deep, whorls flattened laterally above periphery and on
basal margin. Color mainly leached from shell with traces of reddish
flammulations remaining. Umbilicus moderately open, U-shaped,
slightly and regularly decoiling, contained 3.84 times in the diame-
ter. Aperture subcircular, flattened laterally above periphery and on
basal margin, inclined about 20 from the shell axis. Height of lec-
totype 2.47 mm., diameter 4.67 mm.
Lectotype. Cook Islands: Rarotonga. Collected by
Andrew Garrett. ANSP 47754.
Range. One mountain ravine on Rarotonga,
Cook Islands.
Paratypes.BPEM. 2313, ANSP.
Material. Rarotonga (17 specimens, BPBM 2313,
BPBM 167422, ANSP 47754, SMF 165709, FMNH,
NMWC).
Remarks. Although Garrett (1881, p. 387) con-
sidered Sinployea decorticata and S. otareae to be "va-
rieties" of one species, the differences in sculpture and
shape (table XXIII) are sufficient to maintain specific
separation. The two species were restricted to separate
valleys "about three miles apart" (Garrett, 1881, p.
387).
Sinployea harveyensis (Garrett, 1872). Figure 48a-c.
Pitys harveyensis Garrett, 1872, Am. J. Conchol., 7 (4), p. 228, pi.
19, fig. 20 common on Rarotonga, Cook Islands.
Patula harveyensis (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy,
5, p. 93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p.
387.
Helix harveyensis (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p.
542.
Helix (Patula) harveyensis (Garrett), Tryon, 1887, Man. Conchol.,
(2)3, p. 40, pi. 8, figs. 51-53.
Endodonta (Champa) harveyensis (Garrett), Pilsbry, 1893, Man.
Conchol., (2) 9, p. 35.
Diagnosis. Shell very large, diameter 3.91-5.23 mm. (mean
4.35 mm.), with 414-5 normally coiled whorls. Apex and spire rather
strongly elevated, slightly rounded above, body whorl descending
more rapidly, spire protrusion about 14 body whorl width, H/D ratio
0.475-0.641 (mean 0.550). Apical sculpture of fine spiral cords,
eroded or too faint to count accurately. Postnuclear whorls with fine,
irregular, closely spaced, protractively sinuated radial ribs, too
crowded and irregular to count on body whorl, whose interstices are
often less than their width near aperture. Microsculpture visible on
upper spire as very fine radial riblets, 3- 7 between each pair of major
ribs, crossed by slightly finer and more crowded spiral riblets. Um-
bilicus rather narrow, U-shaped, regularly decoiling, contained
3.65-5.48 times (mean 4.73) in the diameter, margins rounded.
Whorls somewhat flattened laterally above periphery, very slightly
on basal margin.
Sinployea harveyensis (fig. 48a-c) is characterized
by its crowded, irregular major radial ribs and elevated
spire. Sinployea youngi (fig. 48d-f) has a wider um-
bilicus, lower spire, more depressed shape, and less
crowded radial ribbing (table XIX).
SYSTEMATIC REVIEW
111
TABLE XXII. - LOCAL VARIATION IN COOK ISLAND SINPLOYEA, II.
.
NUMBER OF
SPECIMENS
RIBS
HEIGHT
DIAMETER
H/D RATIO
planospira
Zurich 3
128.0+20.02
(108-148)
1.77+0.226
(1.52-2.22)
3.7410.250
(3.44-4.24)
0.46810.0277
(0.439-0.524)
youngi
Zurich 3
102.0+3.05
(96-106)
2.0210.119
(1.85-2.25)
4.0610.154
(3.77-4.30)
0.49710.0139
(0.476-0.523)
BPBM 2315 4
127.87.43
(107-139)
2.47+0.098
(2.28-2.68)
4.76+0.117
(4.57-5.10)
0.518+0.0135
(0.493-0.553)
ANSP 47703 3
117.712.33
(114-122)
2.3810.066
(2.32-2.52)
4.9310.077
(4.80-5.07)
0.48310.0156
(0.457-0.510)
harveyensis
ANSP 47756 10
2 19+0 045
4 12+Q 033
531+0 0092
(1.92-2.35)
2 31+0 O''!
(3.91-4.27)
4 21 + 76
(0.475-0.572)
548+0 0065
Zurich 4
(2.25-2.35)
(4.11-4.44)
(0.530-0.560)
BPBM 2343, 8
BPBM 167423
REDUCED
2.6410.077
(2.32-3.01)
4.71+0.111
(4.37-5.23)
0.561+0.0085
(0.530-0.604)
canalis
Zurich, AMS 4
117.318.50
(98-133)
2.0310.068
(1.85-2.15)
4.5210.151
(4.17-4.90)
0.44910.0055
(0.439-0.464)
ANSP 47752 5
122.0112.23
(106-146)
2.0910.055
(1.89-2.19)
4.62+0.156
(4.04-4.90)
0.45210.0103
(0.427-0.481)
BPBM 2328 5
120.8+5.46
(107-130)
2.2810.119
(2.02-2.65)
4.7210.197
(4.40-5.50)
0.48410.0170
(0.446-0.544)
WHORLS
UMBILICUS
D/U RATIO
APICAL CORDS
planospira
Zurich
6 5/8(6 1/4-7 1/4)
1 . 28+0.076
2 90+0 075
(1.15-1.41)
(2.75-2.98)
youngi
Zurich
4+(4-4 1/4)
1.04+0.016
(1.02-1.09)
3.8710.098
(3.68-4.00)
11.3310.34
(11-12)
BPBM
4 l/2-(4 3/8-4 5/8)
1.3410.097
(1.15-1.61)
3.5710.190
(3.15-4.06)
10.2510.25
(10-11)
ANSP
4 3/4+(4 1/2-5)
1.33+0.115
(1.12-1.51)
3.75+0.323
(3.33-4.39)
10.6710.66
(10-12)
harveyensis
4 1 / 2+ (4 3 / 8 4 7 / 8 )
89+0 031
4 61+0 1 36
Zurich
4 1/2(4 I/ 44 5/8)
(0.79-1.12)
86+0 056
(3.65-5.08)
4 89+0 240
(0.76-1.02)
(4.32-5.48)
BPBM
4 7/8-(4 5/8-5 1/8)
0.9910.039
(0.82-1.18)
4.77+0.156
(4.11-5.29)
canalis
Zurich
4 3/4-K4 1/2-5 1/8)
1.5310.064
(1.38-1.68)
2.9310.030
(2.87-3.00)
10.2510.25
(10-11)
ANSP
5-(4 5/8-5 1/4)
1.6310.076
(1.38-1.84)
2.8310.054
(2.64-2.96)
9.60+0.60
(8-11)
BPBM
5+(4 3/4-5 1/2)
1.61+0.079
(1.41-1.84)
2.97+0.067
(2.73-3.10)
10.80+0.49
(9-12)
Description. Shell very large, with 4% normally coiled whorls.
Apex and spire slightly elevated, body whorl descending a little more
rapidly, H/D ratio 0.524. Apical whorls 1%, sculpture mainly eroded
with traces of spiral ribbing remaining. Postnuclear whorls with low,
rounded, irregularly spaced, protractively sinuated radial ribs, be-
coming indistinct on body whorl. Microsculpture a lattice of coequal
radial and spiral riblets. Sutures moderately impressed, whorls flat-
tened laterally above periphery and on basal margin. Color light
yellow-horn with widely spaced, reddish markings absent on base of
shell. Umbilicus U-shaped, relatively narrow, regularly derailing,
contained 4.66 times in the diameter. Aperture ovate, flattened lat-
erally above periphery and slightly on basal margin, inclined about
20 from the shell axis. Height of lectotype 2.17 mm., diameter 4.14
TABLE XXIII. - LOCAL VARIATION IN COOK ISLAND SINPLOYEA, III.
NUMBER OF
SPECIMENS RIBS
HEIGHT
DIAMETER
H/D RATIO
rudis
2.12+0.047
(1.85-2.32)
2.440.088
(2.28-2.85)
3.950.049
(3.71-4.
4.26+0.135
(3.94-4.
0.537+0.0085
30) (0.500-0.575)
0.5740.0165
83) (0.520-0.642)
ANSP 47701 10
BPBM 2320, 6
BPBM 167420
decorticata
89.513.54
(76-103)
2.410.118
(2.19-2.98)
4.290.170
(3.94-5.
0.562+0.0118
10) (0.520-0.590)
BPBM 2313 6
BPBM 47754 4
otareae
73
2.53+0.051
(2.45-2.68)
2.34+0.083
(2.07-2.63)
4.630.079
(4.44-4.
4.570.08A
(4.38-4.
0.5470.0090
80) (0.528-0.566)
0.511+0.0097
90) (0.474-0.540)
BPBM 167418-9 8
BPBM 2335 7
107.05.87
(96-116)
2.540.092
(2.25-2.98)
4.800.097
(4.47-5.
0.5290.0136
23) (0.479-0.570)
ANSP 47755 8
92.293.76
(81-110)
2.450.039
(2.32-2.65)
4.830.065
(4.54-5.
0.507+0.0088
00) (0.470-0.555)
tenulcostata
46.01.00
(45-47)
2.040.083
(1.95-2.12)
4.140.033
(4.11-4.
0.4910.0163
17) (0.475-0.508)
ANSP 47702 2
Zurich 3
47.71.20
(46-50)
2.33+0.040
(2.25-2.38)
4.550.192
(4.20-4.
0.5130.0133
87) (0.489-0.535)
BPBM 2325 5
51.43.65
(43-65)
2.46+0.049
(2.38-2.65)
4.960.171
(4.44-5.
0.4990.0170
46) (0.437-0.537)
WHORLS
UMBILICUS
D/U RATIO
APICAL CORDS
rudis
4 l/4-(3 7/8-4 3/8)
0.800.016
(0.76-0.92)
4.900.111
(4.18-5.42)
10.430.31
(9-12)
ANSP
BPBM
4 3/8+(4 1/4-4 5.8)
0.880.048
(0.79-1.12)
4.830.143
(4.30-5.17)
9.000.58
(8-10)
decort.
4 5/8+(4 3/8-5)
0.900.036
(0.79-1.02)
4.770.257
(4.20-5.92)
8.500.50
(8-9)
BPBM
BPBM
otareae
4 3/4(4 1/2-5)
4 5/8+(4 1/2-5)
1.130.044
(1.02-1.22)
1.11+0.031
(1.02-1.25)
4.06+0.099
(3.84-4.32)
4.110.051
(3.92-4.29)
8
10.000.46
(8-12)
BPBM
BPBM
4 7/8(4 3/4-5)
1.22+0.036
(1.09-1.35)
3.910.061
(3.71-4.22)
9.290.42
(8-11)
ANSP
4 3/4+(4 1/2-5)
1.160.064
(0.89-1.41)
4.240.230
(3.47-5.52)
8.870.40
(8-11)
tenuico.
4(3 7/8-4 1/8)
0.8210.033
(0.79-0.86)
5.010.240
(4.77-5.25)
11.50+0.50
(11-12)
ANSP
Zurich
4 1/8(4-4 1/4)
0.840.040
(0.79-0.92)
5.350.087
(5.25-5.52)
10.000.58
(9-11)
BPBM
4 3/8-(4-4 1/2)
1.000.074
(0.82-1.18)
5.02+0.327
(4.29-6.20)
10.000.32
(9-11)
112
a-f
FIG. 48. a-c, Sinployea harveyensis (Garrett). Rarotonga, Cook Islands. Lectotype. BPBM 2343; d-f, Sinployea youngi (Garrett). Rarotonga,
Cook Islands. Paratype. BPBM 2315. Scale line equals 1 mm. (a-c, MM; d-f, SH).
113
114
SOLEM: ENDODONTOID LAND SNAILS
Lectotype. Cook Islands: Rarotonga. Collected by
Andrew Garrett. ANSP 47756.
Range. Common on Rarotonga, Cook Islands.
Paratypes. BPBM 2343, ANSP.
Material. Rarotonga (20 specimens, BPBM
167423, ANSP 47756, FMNH 46258, Zurich).
Remarks. Garrett did not state whether
Sinployea harveyensis was widely distributed or re-
stricted in its habitat "under rotten wood" (Garrett,
1872, p. 228). Usually he specified if only a single val-
ley was involved. I suspect that S. harveyensis had a
wide range on the island. The very fine radial
sculpture is similar to that seen in the Society Island
Sinployea, but S. harveyensis lacks the prominent
periostracal extensions that characterize the former.
Garrett's retention of large examples is very clearly
shown in Table XXII.
Sinployea youngi (Garrett, 1872). Figure 48d-f.
Pitysyoungi Garrett, 1872, Am. J. Conchol., 7 (4), p. 229, pi. 19, fig.
22 a single valley on Rarotonga, Cook Islands.
Patula youngi (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5, p.
93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 388.
Helix youngi (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 543.
Helix (Patula) youngi (Garrett), Tryon, 1887, Man. Conchol., (2) 3,
p. 40, pi. 8, figs. 54-56.
Endodonta (Champa} youngi (Garrett), Pilsbry, 1893, Man. Con-
chol., (2) 9, p. 35.
Diagnosis. Shell very large, diameter 3.75-5.07 mm. (mean
4.54 mm.), with 3%-5 loosely coiled whorls. Apex and spire moder-
ately and evenly elevated, last whorl descending more rapidly, spire
protrusion about 'A body whorl width, H/D ratio 0.457-0.553 (mean
0.499). Apical sculpture of 10-12 (mean 10.73) prominent spiral
cords. Postnuclear whorls with prominent, large, protractively
sinuated radial ribs, 96-139 (mean 115.4) on the body whorl, whose
interstices are about twice their width. Ribs/mm. 7.15-9.68 (mean
8.13). Microsculpture of fine radial riblets, 3-6 between each pair of
major ribs, crossed by slightly lower and equally crowded spiral rib-
lets. No secondary spiral cording visible. Umbilicus widely open,
V-shaped, last whorl decoiling more rapidly, contained 3.15-4.39
times (mean 3.73) in the diameter, margins rounded. Whorls
strongly flattened laterally above periphery and slightly on basal
margin.
Sinployea youngi (fig. 48d-f) is very similar to S.
harveyensis (fig. 48a-c) but differs in its more de-
pressed shape, lower spire, wider umbilicus, and less
crowded ribbing. Sinployea canalis (fig. 49a-c) has a
much wider umbilicus, much more depressed shape,
greater whorl count, and more laterally compressed
whorl contour. Other species from Rarotonga differ
most obviously in their more widely spaced ribbing or
much smaller size.
Description. Shell very large, with 4% relatively loosely coiled
whorls. Apex and spire slightly and evenly elevated, body whorl
descending more rapidly, H/D ratio 0.510. Apical whorls 1M>,
sculpture of 10 prominent spiral cords, microsculpture obscured.
Postnuclear whorls with prominent, rounded, protractively sinuated
radial ribs, about 117 on the body whorl where they become slightly
irregular, whose interstices are 3-5 times their width. Micro-
sculpture a lattice of very fine radial riblets, 3-6 between each pair
of major ribs, and slightly smaller, more crowded, spiral riblets. Su-
tures moderately impressed, whorls flattened laterally above
periphery and basally. Color dark reddish purple-brown. Umbilicus
V-shaped, moderately open, regularly decoiling, contained 4.39
times in the diameter. Aperture ovate, strongly flattened laterally
above periphery and basally, inclined about 20 from the shell axis.
Height of lectotype 2.50 mm., diameter 4.91 mm.
Lectotype. Cook Islands: Rarotonga. Collected by
Andrew Garrett. ANSP 47703.
Range. One valley on Rarotonga, Cook Islands.
Paratypes. BPBM 2315, ANSP.
Material. Rarotonga (13 specimens, ex BPBM
2339, FMNH 46502, Zurich).
Remarks. Although reported as being collected
"in one valley, the habitat of the typical decorticata"
(Garrett, 1881, p. 388), it was not specified if the
species were sympatric or allopatric. Sinployea decor-
ticata has fewer and much more widely spaced radial
ribs and a more elevated spire than does S. youngi.
Sinployea harveyensis is more similar, but has the api-
cal cords reduced and the radial ribbing quite crowded
and irregular on the body whorl besides differing in
proportions (table XXII).
Sinployea youngi was "A somewhat rare species.
. . . On the ground in damp woods" (Garrett, 1872, p.
229).
Sinployea canalis (Garrett, 1872). Figure 49a-c.
Pitys canalis Garrett, 1872, Am. J. Conchol., 7 (4), p. 227, pi. 19,
fig. 17 one valley on Rarotonga, Cook Islands.
Patula canalis (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5, p.
93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 385.
Helix canalis (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 546.
Helix (Patula) canalis (Garrett), Tryon, 1887, Man. Conchol., (2) 3,
p. 39, pi. 8, figs. 38-40.
Endodonta (Charopa) canalis (Garrett), Pilsbry, 1893, Man. Con-
chol., (2) 9, p. 35.
Diagnosis. Shell very large, diameter 4.01-5.46 mm. (mean
4.63 mm.), with 4M>-5'/2 tightly coiled whorls. Apex and early spire
' fiat or barely elevated, lower spire descending slightly, body whorl
descending sharply, spire protrusion about 'Ao body whorl width, H/D
ratio 0.427-0.544 (mean 0.463). Apical sculpture of 8-12 (mean
10.21) prominent spiral cords. Postnuclear whorls with fine, narrow,
rather closely spaced, protractively sinuated radial ribs, 98-146
(mean 119.8) on the body whorl, whose interstices are 3-4 times
their width. Ribs/mm. 7.07-9.74 (mean 8.22). Microsculpture of
rather widely spaced radial riblets, 3-5 between each pair of major
ribs, crossed by slightly finer and more crowded spiral riblets. Occa-
sionally weak secondary spiral cording visible in umbilicus and su-
tures. Umbilicus broadly open, cup-shaped, regularly decoiling, con-
tained 2.64-3.10 times (mean 2.91) in the diameter. Sutures very
deep, whorls shouldered above, nearly vertically compressed on outer
margin, basal margin strongly rounded, aperture inclined about 25
from shell axis.
The widely open umbilicus, strongly laterally
compressed whorls, fine ribbing, and large size char-
acterize Sinployea canalis (fig. 49a-c). Other Cook Is-
land species of similar size have much more widely
spaced ribbing and narrower umbilici (S. otareae, fig.
50a-c, and S. tenuicostata, fig. 50d-f) or normally con-
toured whorls and a narrower umbilicus (S. youngi, fig.
48d-f). The slightly larger Samoan S. complementaria
(fig. 56a-f) has a much narrower umbilicus, fewer and
more widely spaced radial ribs, and many more micro-
radial riblets.
Description. Shell very large, with slightly less than 5!4 rather
tightly coiled whorls. Apex and spire slightly sunken, body whorl
SYSTEMATIC REVIEW
115
FIG. 49. a-c, Sinployea canalis (Garrett). Rarotonga, Cook Is-
lands. Paratype. BPBM 2328. Scale line equals 2 mm. (SH).
descending moderately, H/D ratio 0.427. Apical whorls 1%, sculpture
of 8 prominent spiral ribs with a barely visible radial microsculpture.
Postnuclear whorls with low, rounded, relatively crowded, protrac-
tively sinuated radial ribs, 106 on the body whorl, whose interstices
are 2-3 times their width. Microsculpture a lattice of fine radial
riblets, 3-5 between each pair of major ribs, crossed by slightly finer
and more crowded spiral riblets with a secondary sculpture of spiral
cording visible in suture and umbilicus. Sutures deep, whorls shoul-
dered above, flattened laterally above periphery and on basal mar-
gin. Color yellowish horn with extensive vague reddish flammula-
tions. Umbilicus broadly open, cup-shaped, regularly decoiling,
contained 2.96 times in the diameter. Aperture subovate, strongly
flattened laterally above periphery and on basal margin, inclined
about 25 from the shell axis. Height of lectotype 2.07 mm., diameter
4.77 mm.
Lectotype. Cook Islands: Rarotonga. Collected by
Andrew Garrett. ANSP 47752.
Range. One valley on Rarotonga, Cook Islands.
Paratypes.BPBM 2328, ANSP.
Material. Rarotonga (13 specimens, AMS,
Zurich, SMF 165456).
Remarks. The strong lateral compression of the
body whorl is not duplicated by any of the other
Polynesian species and provides a strong contrast (fig.
49b) to the other Rarotongan species. Garrett (1872, p.
227) recorded this as a "somewhat rare species, found
on the ground in damp forests, and confined to a single
valley." Measured sets (table XXII) show very little
variation and suggest fractions of a single population.
Sinployea otareae (Garrett, 1872). Figure 50a-c.
Pitys otareae Garrett, 1872, Am. J. Conchol., 7 (4), pp. 228-229, pi.
19, fig. 21 one valley on Rarotonga, Cook Islands.
Patula otareae (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5, p.
94.
Helix otareae (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 543.
Patula decorticata var. otareae (Garrett), Garrett, 1881, J. Acad.
Nat. Sci., Philadelphia, 8 (4), p. 387.
Helix (Patula) decorticata var. otareae (Garrett), Tryon, 1887, Man.
Conchol., (2) 3, p. 40, pi. 8, fig. 50.
Endodonta (Charopa) decorticata var. otareae (Garrett), Pilsbry,
1893, Man. Conchol., (2) 9, p. 35.
Diagnosis. Shell very large, diameter 4.21-5.20 mm. (mean
4.68 mm.), with 4'/2-5 relatively tightly coiled whorls. Apex and
spire moderately and evenly elevated, last whorl descending more
rapidly, spire protrusion more than *k body whorl width, H/D ratio
0.470-0.570 (mean 0.516). Apical sculpture of 8-12 (mean 9.41)
prominent spiral cords. Postnuclear whorls, with narrow, prominent,
protractively sinuated, rather widely spaced radial ribs, 81-116
(mean 95.6) on the body whorl, whose interstices are 2-3 times their
width. Ribs/mm. 5.72-7.73 (mean 6.48). Microsculpture of fine,
crowded radial riblets, 6-10 between each pair of major ribs, crossed
by finer and more crowded spiral riblets. No secondary spiral cord-
ing. Umbilicus open, V-shaped, regularly decoiling, contained 3.47-
5.52 times (mean 4.10) in the diameter, margins rounded. Whorl
contours and aperture typical.
The presence of only six to ten microradials be-
tween each pair of major ribs at once separates
Sinployea otareae (fig. 50a-c) from S. decorticata (fig.
47a-c). Individual specimens may show the size and
shape characters of the other species, but the sculp-
tural difference is consistent. Sinployea tenuicostata
(fig. 50d-f) has a flat spire, fewer ribs, and a narrower
umbilicus.
FIG. 50. a-c, Sinployea otareae (Garrett). Rarotonga, Cook Islands. Paratype. BPBM 2335; d-f, Sinployea tenuicostata (Garrett). Rarotonga,
Cook Islands. Paratype. BPBM 2325. Scale lines equal 1 mm. (o-c, MM; d-f, SH).
116
SYSTEMATIC REVIEW
117
Description. Shell extremely large, with 4% relatively tightly
coiled whorls. Apex and early spire flat, later whorls descending
gradually, body whorl descending moderately, H/D ratio 0.470. Api-
cal whorls 1%, sculpture of 8 prominent spiral ribs with a micro-
sculpture of radial corrugations. Postnuclear whorls with prominent,
high, protractively sinuated radial ribs, 110 on the body whorl,
whose interstices are 2-3 times their width. Microsculpture a lattice
of radial riblets, 6-9 between each pair of major ribs, and slightly
smaller, more crowded spiral riblets. Sutures deep, whorls flattened
laterally above periphery and on basal margin. Color reddish
yellow-brown. Umbilicus moderately open, V-shaped, regularly de-
coiling, contained 3.98 times in the diameter. Aperture ovate, flat-
tened laterally above periphery and basally, inclined about 20 from
the shell axis. Height of lectotype 2.34 mm., diameter 4.97 mm.
Lectotype. Cook Islands: Rarotonga. Collected by
Andrew Garrett. ANSP 47755.
Range. Under dead wood on the banks of a
stream in a single valley on Rarotonga, Cook Islands.
Paratypes. BPBM 2335, ANSP.
Material. Cook Islands (2 specimens, NMWC):
Rarotonga (30 specimens, BPBM 167418-9, FMNH
46411, FMNH 90615).
Remarks. Mautodontha zebrina came from the
same unidentified valley (Garrett, 1881, p. 387).
Sinployea otareae was abundant and "Living examples
are uniform deep black" (Garrett, 1872, p. 229). Only
minor size variation was observed (table XXIII).
Sinployea tenuicostata (Garrett, 1872). Figure
50d-f.
Pitys tenuicostata Garrett, 1872, Am. J. Conchol., 7 (4), p. 229, pi.
19, fig. 23 rare, but widely distributed on Rarotonga, Cook
Islands.
Patula tenuicostata (Garrett), Schmeltz, 1874, Cat. Mus. Godeftroy,
5, p. 94; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p.
385.
Helix tenuicostata (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p.
547.
Helix (Patula) tenuicostata (Garrett), Tryon, 1887, Man. Conchol.,
(2)3, p. 39, pi. 8, figs. 35-37.
Endodonta (Champa) tenuicostata (Garrett), Pilsbry, 1893, Man.
Conchol., (2) 9, p. 35.
Diagnosis. Shell very large, diameter 4.08-5.43 mm. (mean
4.70 mm.), with 3%-4'/4 relatively loosely coiled whorls. Apex and
early spire flat or barely depressed, later whorls descending slightly,
H/D ratio 0.437-0.537 (mean 0.499). Apical sculpture of 9-12 (mean
10.45) prominent spiral cords. Postnuclear whorls with narrow,
widely spaced, protractively sinuated radial ribs, 43-66 (mean 50.7)
on the body whorl, whose interstices are 3-6 times their width.
Ribs/mm. 2.87-4.23 (mean 3.46). Microsculpture of rather large and
widely spaced radial riblets, 8-12 between each pair of major ribs,
crossed by much finer and more crowded spiral riblets. No secondary
spiral cording. Umbilicus relatively narrow, U-shaped, regularly de-
coiling, contained 4.29-6.20 times (mean 5.16) in the diameter, mar-
gins rounded. Aperture strongly flattened laterally above periphery.
Sinployea tenuicostata (fig. 50d-f) has few and
widely spaced major radial ribs, a flat spire, and rela-
tively few and larger microradial riblets. The Samoan
S. complementaria (fig. 56a-f) has an elevated spire,
more major radials, and much finer and more numer-
ous microradial riblets. Sinployea otareae (fig. 50a-c) is
the same size, but has an elevated spire, many more
radial ribs, and a wider umbilicus.
Description. Shell very large, with 4Vs loosely coiled whorls.
Apex and spire fiat, last whorl descending slightly, H/D ratio 0.508.
Apical whorls 1%, sculpture of 11 moderately prominent spiral ribs
with a faint microsculpture of radial corrugations. Postnuclear
whorls with narrow, prominent, rather widely spaced, protractively
sinuated radial ribs, 47 on the body whorl, whose interstices are 3-5
times their width. Microsculpture of fine, rather widely spaced radial
riblets, 6-10 between each pair of major ribs, crossed by much finer
and more crowded spiral riblets. Sutures deep, whorls strongly
rounded above, flattened laterally above periphery and slightly on
basal margin. Color light yellow-horn without darker maculations.
Umbilicus narrow, U-shaped, regularly decoiling, contained 5.25
times in the diameter. Aperture subcircular, flattened laterally
above periphery, inclined about 25 from the shell axis. Height of
lectotype 2.11 mm., diameter 4.15 mm.
Lectotype. Cook Islands: Rarotonga. Collected by
Andrew Garrett. ANSP 47702.
Range. Widely distributed on Rarotonga, Cook
Islands.
Paratypes. BPBM 2325, ANSP.
Material. Rarotonga (8 specimens, Zurich, SMF
165748).
Remarks. Originally described as a "somewhat
rare species, having a wide range on the island, and
generally found on the ground on the sides of ravines"
(Garrett, 1872, p. 229), later Sinployea tenuicostata
was termed "comparatively rare" (Garrett, 1881, p.
385). Available sets showed considerable size variation
(table XXIII), which is probably magnified by the lack
of clear differentiation into preadult and gerontic
growth.
SAMOAN AND SWAINS ISLAND Sinployea
Except for Ofu and Olosega, every island in the
Samoan group that has been collected on has yielded at
least one species of Sinployea. Only one endodontid,
Minidonta manuaensis Solem (1976b, p. 130), has been
collected on Olosega, and no specimens of this family
have been taken on Ofu. Although only two examples
of Sinployea clista have been taken on Tutuila, com-
paratively little collecting has been done on that is-
land. Aunuu Island off Tutuila has an endemic species,
S. aunuuana, which also is recorded from Ta'u on the
basis of a single very worn individual that may repre-
sent an undescribed species. Sinployea clausa and S.
allecta tauensis are restricted to Ta'u in the Manu'a
Group. All three Ta'u species have been found at a
single station (Station 76), and S. clausa and S. a.
tauensis were collected together at Stations 75, 184,
188, and 189. None of the Ta'u examples were collected
alive. Scattered collections from Savai'i produced
abundant material of Sinployea allecta allecta (Cox)
from lowland localities. Much less abundant material
of a more heavily ribbed form of this subspecies was
collected from diverse localities and at several eleva-
tions on Upolu. At the foot of Mt. Tafua Upolu (Station
7), S. clista was taken together with S. a. allecta,
whereas both S. clista and S. complementaria (Mous-
son) occurred at the foot of Mt. Solaua (Station 18). The
latter is the dominant species on Upolu, and I suspect it
is significant that S. allecta (Cox) and S. complemen-
taria (Mousson) have not been collected at the same
118
SOLEM: ENDODONTOID LAND SNAILS
stations. Swains Island has an endemic species, Sin-
ployea intermedia.
No adult examples of Upolu S. allecta allecta were
available for dissection, so the degree of difference from
S. clista remains uncertain. The latter has a penis
about 1.2 mm. long, whereas Savai'i examples of S.
allecta have a 1.3-1.5 mm. penis with vergic papilla
enlarged and stimulatory pad reduced in size. Sin-
ployea complementaria is immediately differentiated
by its very large penis of more than 2.0 mm. in length.
Study of Upolu S. allecta would be desirable in order to
determine the pattern of species isolation in penis
structure.
Compared with the Cook and Society Island
species, Samoan Sinployea tend to have many radial
microriblets, lack secondary spiral cording (except S.
clausa and S. clista), tend toward a narrowed or closed
umbilicus (except S. aunuuana), and have a flat apex
but elevated spire (table XXIV). Sinployea aunuuana
(fig. 52a-c) has the widest umbilicus and relatively
crowded major radial ribbing. Its flat spire and more
rounded body whorl ally it to the Samoan taxa.
Sinployea clista (fig. 51a-c) and S. clausa (fig. 51d-f)
are small species with rather widely spaced ribbing
whose umbilici are closed or with only a narrow lateral
crack. Sinployea clausa has the umbilicus open to the
apex and the spire more elevated than in S. clista
where the umbilicus is never open to the apex. Both of
these have rather prominent secondary spiral cording.
Sinployea allecta (fig. 54a-c) and S. intermedia (fig.
52d-f) are larger, have a more open umbilicus, at most
faint traces of secondary spiral cording, and rather
widely spaced major radial ribs. Sinployea com-
plementaria (fig. 56a-f) is the largest known species in
the genus, has an increased whorl count, very few ra-
dial ribs, and a moderately open umbilicus. Sinployea
intermedia is the easternmost Polynesian species to
lack gray pigmentation on the head and ommato-
phores. Since Swains Island is low and has coastal
scrub rather than dense forest, restriction of this
species to the ground stratum and loss of pigmentation
would not be surprising.
TABLE XXIV. - RANGE OF VARIATION IN SAMOAN AND SWAINS ISLAND SINPLOYEA.
NUMBER OF
SPECIMENS
clausa
36
10
136
clista
aunuuana
allecta allecta
(Cox)
ALL
151
Savaii
86
Upolu
65
allecta tauensis
26
46
126
intermedia
complement aria
(Mo us son)
RIBS
RIBS/MM.
HEIGHT
DIAMETER
H/D RATIO
86.6(67-106) 11.80(9.04-14.23) 1.51(1.25-1.97) 2.41(2.11-2.93) 0.625(0.577-0.732)
98.3(74-115) 11.98(9.13-14.36) 1.50(1.28-1.84) 2.53(2.17-2.96) 0.591(0.555-0.636)
112.6(92-131) 13.13(11.18-14.57) 1.40(1.25-1.71) 2.76(2.57-3.09) 0.509(0.465-0.572)
89.1(56-121) 9.00(5.83-11.98) 1.77(1.51-2.04) 3.13(2.63-3.62) 0.571(0.511-0.622)
75.9(56-89) 7.84(5.85-8.97) 1.75(1.51-1.94) 3.07(2.65-3.36) 0.576(0.558-0.598)
99.3(84-121) 9.90(8.48-11.98) 1.79(1.51-2.04)
79.8(61-101) 8.31(6.20-10.51) 1.73(1.61-1.97)
82.1(63-96) 7.66(5.75-9.16) 2.03(1.82-2.19)
69.1(52-95) 4.50(3.11-5.79) 2.55(2.11-3.32)
3.17(2.63-3.62) 0.567(0.511-0.622)
3.00(2.70-3.39) 0.577(0.542-0.616)
3.49(3.21-3.81) 0.581(0.534-0.616)
4.92(4.31-5.92) 0.517(0.458-0.648)
aunuu.
a. all.
ALL
Sav.
Upo.
compl .
WHORLS
UMBILICUS
D/U RATIO
APICAL
CORDS
SPIRE
ELEVATION
BODY WHORL
WIDTH
SP/BWW
claus - 3 7/8-(3 1/2-4 3/8) 0.11(0.07-0.20) 23.1(13.7-39) 9.40(8-10) 0.19(0.16-0.23) 0.90(0.82-0.92) 0.215(0.200-0.233)
cllst. 3 3/4+(3 1/2-4 1/4) LESS THAN 0.13 ^QW"^ 9 - 17 <S-U> 0.07(0.03-0.13) 0.98(0.86-1.12) 0.089(0.033-0.176)
4 l/8-(4-4 1/2) 0.75(0.66-0.95) 3.69(3.24-4.05) 9.50(8-12) 0.13(0.07-0.26) 0.89(0.82-0.92) 0.125(0.074-0.286)
4 1/8(3 5/8-4 1/2) 0.41(0.26-0.59) 7.85(5.93-9.80) 8.60(7-10) 0.14(0.07-0.23) 1.11(0.99-1.22) 0.123(0.056-0.250)
4+(3 5/8-4 3/8) 0.39(0.33-0.43) 7.89(6.85-9.20) 8.50(8-9) 0.16(0.12-0.21) 1.09(0.99-1.15) 0.143(0.103-0.191)
4 1/8+0 7/8-4 1/2) 0.42(0.26-0.59) 7.82(5.93-9.80) 8.70(7-10) 0.13(0.07-0.23) 1.12(1.02-1.22) 0.117(0.056-0.250)
4-(3 3/4-4 1/8) 0.46(0.33-0.69) 6.80(4.72-9.55) UNKNOWN 0.19(0.18-0.20) 1.09(1.05-1.12) 0.172(0.162-0.188)
4 3/8+(4 1/4-4 3/4) 0.64(0.53-0.79) 5.43(4.59-6.12) 10.57(9-13) 0.23(0.13-0.35) 1.12(1.02-1.25) 0.209(0.125-0.309)
4 5/8+(4 3/8-5 1/4) 0.99(0.76-1.22) 4.99(4.18-6.08) 10.08(8-13) 0.22(0.07-0.46) 1.55(1.35-1.81) 0.140(0.039-0.275)
SYSTEMATIC REVIEW
119
KEY TO SAMOAN AND SWAINS ISLAND Sinployea
1. Umbilicus closed or D/U ratio averaging more than 20 2
Umbilicus open, D/U ratio averaging much less than 10 3
2. Spire protrusion about Vs body whorl width; umbilicus with
apex visible Sinployea clausa, new species
Spire protrusion averaging less than 0.10; umbilicus closed or
at most a lateral crack Sinployea clista, new species
3. Diameter less than 4.00 mm.; ribs/mm, averaging more than
6.00 4
Diameter more than 4.00 mm.; ribs/mm, averaging less than
5.00 Sinployea complementaria (Mousson, 1865)
4. D/U ratio averaging more than 5.00; H/D ratio averaging more
than 0.550 5
D/U ratio averaging about 3.75; H/D ratio about 0.510
Sinployea aunuuana, new species
5. Mean diameter about 3.0-3.2 mm.; mean D/U ratio more than
6.50 6
Mean diameter about 3.5 mm.; mean D/U ratio about 5.50;
Swains Island Sinployea intermedia, new species
6. Spire more protruded; umbilicus slightly wider; Ta'u, Manu'a
Group Sinployea allecta tauensis, new subspecies
Spire less protruded; umbilicus slightly narrower; Upolu and
Savai'i Sinployea allecta allecta (Cox, 1870)
Sinployea clausa, new species. Figure 51d-f.
Diagnosis. Shell small, diameter 2.11-2.93 mm. (mean 2.41
mm.), with 3'A-4 3 /8 normally coiled whorls. Apex and spire moder-
ately and evenly elevated, body whorl descending much more
rapidly, spire protrusion more than 'k body whorl width, H/D ratio
0.577-0.732 (mean 0.625). Apical sculpture of 8-10 (mean 9.40) fine
spiral cords. Postnuclear whorls with narrow, rounded, protractive,
rather inconspicuous radial ribs, 67-106 (mean 86.6) on the body
whorl, whose interstices are 2-4 times their width. Ribs/mm. 9.04-
14.2 (mean 11.80). Microsculpture of extremely fine radial riblets,
5- 10 between each pair of major ribs, crossed by slightly finer and
more crowded spiral riblets. Secondary sculpture of fine, rather
widely spaced spiral cords visible over entire shell surface. Um-
bilicus very narrow, open to apex, about 'A constricted by basal lip,
contained 13.7-39 times (mean 23.1) in the diameter, margins
rounded. Whorl profile almost circular, only flattened slightly on
upper palatal wall, basal margin rounded, aperture inclined about 5
from shell axis.
Sinployea clista (compare fig. 51a-c) is very closely
related, differing primarily in its lower spire and gen-
erally imperforate or at most very narrowly open um-
bilicus. Other Samoan species have wider umbilici. Ex-
tralimital species with narrow umbilici, such as Ba
humbugi (fig. 74a-c) and Sinployea adposita (fig.
69d-f) from Fiji, have vastly different shapes and
sculpture.
Description. Shell small, with 4 tightly coiled whorls. Apex
and spire moderately and evenly elevated, last whorl descending
much more rapidly, H/D ratio 0.613. Embryonic whorls l'/2, sculpture
partly eroded, remnants of about 9 spiral cords. Remaining whorls
with moderately prominent, rounded, protractive radial ribs, 89 on
the body whorl, whose interstices are 2-4 times their width. Micro-
sculpture of very fine radial riblets, 5-12 between each pair of major
ribs, even finer spiral riblets and larger, relatively widely spaced,
secondary spiral cords. Sutures deeply impressed, whorls strongly
rounded above with only slight lateral flattening. Umbilicus barely
perforate, partially constricted by basal lip, contained 15 times in the
diameter. Color a uniform, very light yellow-brown. Aperture circu-
lar with slightly expanded baso-columellar margin, slightly flat-
tened laterally above periphery, inclined about 5 from the shell axis.
Height of holotype 1.51 mm., diameter 2.47 mm.
Holotype. Samoa: Ta'u, Station 188, Faleasao,
700 ft. inland, at 12 ft. elevation. Collected by Wray
Harris on September 24, 1937. BPBM 187736.
Range. Ta'u, Manu'a Group, American Samoa.
Paratypes. Ta'u: Siufaga (Stations 16, 76, 138,
184, 185, 186, 211) at 350-600 ft. inland at 5-500 ft.
elevation (19 specimens, BPBM 171136, BPBM
173784, ex BPBM 171073, ex BPBM 182410, ex BPBM
187653, BPBM 186682, BPBM 187625, BPBM 187658,
BPBM 187705, BPBM 188085); Tavalagi, Vi2 mile in-
land (Station 75) at 500 ft. elevation above Faleasao
cliffs (3 specimens, BPBM 171065, ex BPBM 171106,
ex BPBM 182095); Faleasao, 600-700 ft. inland (Sta-
tions 185, 188-190) at 12 ft. elevations (11 specimens,
BPBM 187665, BPBM 187728, BPBM 187736, BPBM
187774, ex BPBM 187753, ex BPBM 187781, BPBM
187982, BPBM 188277); Amouli, l /z mile inland (Sta-
tion 80) at 500 ft. elevation (1 specimen, BPBM
182150); Palapala, Vfe mile inland (Station 201) at 600
ft. elevation (1 specimen, BPBM 187919).
Remarks. The only available specimens were
dead shells taken from sweepings mixed in coral rub-
ble. A high percentage of the shells had bits of coral
stone and sand cemented to the shell surface, and
many were quite worn. Specimens from Faleasao did
not differ significantly from those taken at Siufaga
(table XXV).
The name clausa, from the Latin meaning closed,
is perhaps slightly misleading, because the umbilicus
is always minutely perforate, but the name was chosen
to emphasize its close relationship to Sinployea clista
from Upolu and Tutuila.
In combining a higher spire and wider umbilicus
than is found inS. clista, S. clausa parallels (fig. 51a-f)
the divergence seen between the subspecies of S. al-
lecta (fig. 54a-e). Only the proportionately greater size
of the morphologic gap between S. clista and S. clausa
accounts for the arbitrary consideration of them as
species rather than subspecies.
Sinployea clista, new species. Figures 51a-c, 53d-e.
Diagnosis. Shell small, diameter 2.17-2.96 mm. (mean 2.53
mm.), with 3 l /2-4'/t normally coiled whorls. Apex and spire flat to
slightly and evenly elevated, body whorl descending much more
rapidly, spire protrusion less than Vio body whorl width, H/D ratio
0.555-0.636 (mean 0.591). Apical sculpture of 8-11 (mean 9.17) nar-
row spiral cords. Postnuclear whorls with low, rounded, protractively
sinuated radial ribs, 74-115 (mean 98.3) on the body whorl, whose
interstices are about 3-4 times their width except becoming crowded
and indistinct on end of body whorl. Ribs/mm. 9.13-14.36 (mean
11.98). Microsculpture of very fine radial riblets, 5-12 between each
pair of major ribs, crossed by slightly finer spiral riblets, relatively
crowded and inconspicuous secondary spiral cording visible on most
of shell surface. Umbilicus either closed (50%) or with a narrow lat-
eral opening that prevents seeing the apex, contained more than 20
times in the diameter. Whorls strongly rounded on all margins, very
slightly flattened laterally above periphery, aperture inclined about
15 from shell axis.
Sinployea clausa (fig. 51d-f) has the umbilicus
narrowly open and the spire distinctly more elevated.
Other minor differences are discussed below. Ba hum-
bugi (fig. 74a-c) from Fiji has a much more elevated
spire (mean H/D ratio 0.817) and very few and widely
spaced major radial ribs. All other Sinployea are at
least narrowly umbilicated.
FIG. 51. a-c, Sinployea clista, new species. Olomoana, Tutuila, Samoa. Holotype. BPBM 84372; d-f, Sinployea clausa, new species. Station
188, Faleasao, Ta'u, Samoa. Holotype. BPBM 187736. Scale lines equal 1 mm. (a-c, SG; d-f, MM).
120
TABLE XXV. - LOCAL VARIATION IN SAMOAN SINPLOYEA, I.
NUMBER OF
SPECIMENS
RIBS
HEIGHT
DIAMETER
H/D RATIO
clausa
Faleasao
Siufaga
10 88.115.88 I.470.063 2.3810.080 0.6150.0006
(75-101) (1.22-1.81) (2.07-2.93) (0.577-0.666)
11 85.614.33 1.5410.050 2.4210.050 0.63710.0122
(67-106) (1.38-1.97) (2.20-2.70) (0.584-0.732)
clista
Tutulla
BPBM 84372
Upolu
BPBM 185879,
BPBM 186642
FMNH 153177,
BMW MF3992
89.016.02
(83-95)
74
112.712.34
(108-115)
1.84
2.9310.033 0.62910.0070
(2.89-2.96) (0.622-0.636)
1.2910.011 2.2310.039 0.53210.0055
(1.28-1.32) (2.17-2.30) (0.572-0.591)
1.4810.050 2.5710.025 0.57510.0130
(1.41-1.58) (2.53-2.63) (0.555-0.600)
aunuuana
BPBM 83242
25
1.4110.018 2.7610.024 0.50810.0050
(1.26-1.72) (2.58-3.11) (0.465-0.556)
allecta allecta
Upolu
AMS, BPBM
Zurich
S avail
Sta. 38
FMNH 153371
18 97.115.10 1.76+0.043 3.0910.064 0.56710.0073
(78-121) (1.39-2.02) (2.52-3.41) (0.511-0.622)
13 101.83.72 1.8410.020 3.2610.044 0.56610.0045
(84-121) (1.71-2.01) (3.03-3.55) (0.543-0.598)
6 73. 411. 86 1.7510.064 3.0410.096 0.5760.0064
(69-80) (1.51-1.94) (2.66-3.36) (0.558-0.598)
WHORLS
UMBILICUS
D/U RATIO
APICAL CORDS
clausa
Fale.
Siuf.
clista
Tutu.
Upolu
BPBM
FMNH
3 3/4+(3 1/2-4 1/8) 0.1210.011
(0.07-0.20)
3 7/8+(3 1/2-4 3/8) 0.1210.012
(0.07-0.20)
4 1/4
0.13
3 5/8(3 1/2-3 3/4) 0.0810.011
(0.07-0.10)
3 5/8+(3 5/8-3 3/4)
22.412.19
(14.8-39)
22.712.07
(13.6-35)
22.5
9.2510.48
(8-10)
10. Oil.
(9-11)
29.913.3 9
(23.3-33.5)
8.510.50
(8-9)
aunuuana
BPBM 4 l/8-(4-4 1/2)
a. allecta
Upolu
AMS 4 1/8(3 7/8-4 3/8)
Zurich 4 l/8+(4-4 3/8)
Savaii
Sta. 38 4-(3 5/8-4 1/4)
0.7410.014
(0.66-0.95)
0.3910.015
(0.26-0.49)
0.4310.020
(0.33-0.53)
0.3910.020
(0.33-0.43)
3.6910.044 9.0010.55
(3.24-4.05) (8-11)
8.0510.250
(5.93-9.70)
7.7410.316
(6.19-9.80)
7.8910.356
(6.85-9.20)
8.3310.67
(7-9)
8.8310.48
(7-10)
(8-9)
121
122
SOLEM: ENDODONTOID LAND SNAILS
Description. Shell small, with 4'/4 normally coiled whorls. Apex
and spire slightly and evenly elevated, last whorl descending much
more rapidly, H/D ratio 0.622. Embryonic whorls IVz, sculpture of 11
narrow spiral cords with an intrusion of vague radial corrugations on
last quarter whorl. Postnuclear whorls with low, rounded, protrac-
tively sinuated radial ribs, 93 on the body whorl, whose interstices
are 2-4 times their width, becoming irregular and more crowded on
gerontic portions of body whorl. Microsculpture of fine radial riblets,
5-10 between each pair of major ribs, exceedingly fine spiral riblets,
and inconspicuous secondary spiral cords. Sutures deep, whorls flatly
rounded above periphery with broadly rounded basal margin, sloping
to narrow umbilical chink. Umbilicus closed by expansion of inner
portion of body whorl and reflection of umbilical lip. Color light
yellow-brown with reddish markings. Aperture circular with slightly
less sharply rounded supraperipheral outer margin, inclined about
15 from shell axis. Height of holotype 1.84 mm., diameter 2.96 mm.
Holotype. Samoa: Tutuila, Olomoana. Collected
under stones and on dead leaves by members of the
Samoan Expedition on March 17, 1926. BPBM 84372.
Range. Tutuila, American Samoa, and Upolu,
Western Samoa.
Paratypes. Tutuila: Olomoana, under stones and
dead leaves (1 specimen, BPBM 84372). Upolu: Mt.
Vaea (Station 147) at 850 ft. elevation (3 specimens,
BPBM 185879); Tatapao-Lake Lanuto'o trail at 1,250
ft. elevation (1 specimen, BPBM 186642); foot of south-
east peak, Mt. Tafua Upolu (Station 7) at 1,200 ft. ele-
vation (1 specimen, FMNH 153081); foot of Mt. Solaua
(Station 18) at 600-800 ft. elevation (2 specimens,
FMNH 153177); Tanumalala at 800 ft. elevation (1
specimen, BMW MF 3992, collected by R. A. Cumber
on May 15, 1954).
Remarks. Besides the obvious differences in
spire protrusion and umbilical closure, Sinployea clista
differs from S. clausa in having the aperture more
strongly inclined and the major ribbing protractively
sinuated rather than being just slightly inclined. The
latter also has the secondary spiral cording more prom-
inent and more widely spaced.
Sinployea clista is sparsely distributed on both
Upolu and Tutuila. Although there are obvious size,
shape, and ribbing differences (tables XXIV- XXV), so
little material is available that I give no systematic
recognition to these variations. Despite intensive ef-
forts after field recognition, no additional examples
could be located at the Mt. Tafua Upolu and Mt. Solaua
stations.
The name clista is taken from the Greek for closed
and refers to the unusual umbilicus.
Description of soft parts. Pallial region (fig. 53e) about 3.5 mm.
long, rectal kidney lobe 1.7 mm. long, much longer than pericardia!
arm. Space between ureter arms wider than in most Polynesian
species.
Genitalia (fig. 53d) fragmentary, observed portions without un-
usual features. Penis about 1.2 mm. long.
(Based on BPBM 84372, 1 partial example.)
Sinployea aunuuana, new species. Figures 52a-c,
53a-c.
Diagnosis. Shell slightly smaller than average, diameter
2.57-3.09 mm. (mean 2.76 mm.), with 4-4'/2 normally coiled whorls.
Apex and early spire flat or slightly protruding, lower spire whorls
descending slightly, body whorl descending rapidly, spire protrusion
about Vx body whorl width, H/D ratio 0.465-0.572 (mean 0.509). Api-
cal sculpture of 8-12 (mean 9.50) narrow spiral cords. Postnuclear
sculpture of very fine, narrow, rather crowded, slightly protractively
sinuated radial ribs, 92-131 (mean 112.6) on the body whorl, whose
interstices are 2-3 times their width. Ribs/mm. 11.18-14.57 (mean
13.13). Microsculpture of very fine radial riblets, 5-9 between each
pair of major ribs, crossed by very fine and crowded spiral riblets,
with an indistinct development of secondary spiral cording occasion-
ally visible. Umbilicus broadly open, V-shaped, regularly decoiling,
contained 3.24-4.05 times (mean 3.69) in the diameter, margins
rounded. Whorl contours slightly less compressed than average,
aperture inclined about 10 from shell axis.
The generally flat apex and early spire of
Sinployea aunuuana (fig. 52a-c) combine with the
wide umbilicus and crowded radial ribbing to easily
separate it from the other Polynesian species. Of
similar-sized species, S. neglecta (fig. 41d-f) from
Huahine has a much more elevated spire, proportion-
ately narrower umbilicus, and about 12.5 apical cords;
S. atiensis (fig. 44d-f) has a higher spire and narrower
umbilicus with broader and more widely spaced ribs.
Sinployea kusaieana (fig. 63a-c) has a narrower body
whorl, fewer whorls, and a proportionately smaller
apex.
Description. Shell slightly smaller than average, with 4'/s
tightly coiled whorls. Apex and early spire flat, body whorl descend-
ing rapidly, spire protrusion about 'Ao of body whorl width, H/D ratio
0.524. Embryonic whorls 1%, sculpture of 12 moderately prominent
spiral ribs. Postnuclear whorls with somewhat irregularly spaced,
high, rounded radial ribs, 123 on the body whorl, whose interstices
are usually about twice their width. Microsculpture of very fine ra-
dial riblets, 3-6 between each pair of major ribs, crossed by barely
visible, spiral riblets that are very crowded. Occasional traces of
weak spiral cording. Sutures deeply impressed, whorls strongly
rounded above, flattened laterally above periphery and slightly on
basal margin. Umbilicus broadly V-shaped, widely open, regularly
decoiling, contained 3.82 times in the diameter. Color uniform light
reddish brown. Aperture circular, strongly flattened laterally above
periphery, inclined about 10 from the shell axis. Height of holotype
1.45 mm., diameter 2.77 mm.
Holotype. Samoa: Aunuu Island (off Tutuila),
Station 14, 500 yd. from shore at 200 ft. elevation. Col-
lected by T. T. Dranga on February 4, 1926. BPBM
83256.
Range. Aunuu Island off Tutuila and probably
Ta'u, Manu'a Group, American Samoa.
Paratypes. Aunuu: 10 yd. east of lighthouse, 200
yd. inland at 250 ft. elevation (134 specimens, BPBM
83242). Ta'u: Siufaga swamp, 400 ft. inland (Station
76) at 7 ft. elevation (1 specimen, ex BPBM 182444).
Remarks. Although the size and umbilical shape
of Sinployea aunuuana (fig. 52a-c) is more similar to
the Cook and Society Island species, the very fine ra-
dial riblets, wide body whorl, flat spire, and round
whorl contours relate it to the other Samoan species.
Sinployea allecta (fig. 54a-e) is most similar, but
differs in its much narrower umbilicus, more elevated
shape (table XXIV), and absence of secondary spiral
cording. They agree in their spire shape and relative
protrusion.
A single dead, very worn specimen from Ta'u is
referred here with some hesitation. In size and proper-
abc
FIG. 52. a-c, Sinployea aunuuana, new species. Station 14, Aunuu Island, Tutuila, Samoa. Holotype. BPBM 83256; d-f, Sinployea inter-
media, new species. 200 yd. inland, west side Swains Island. Holotype. BPBM 186666. Scale lines equal 1 mm. (MM).
123
124
SOLEM: ENDODONTOID LAND SNAILS
-VD
FIG. 53. Anatomy of Sinp/oyea aunuuana, S. clista, and S. intermedia from Samoa and Swains Island: a-c, Sinployea aiinuuana, new
species. Station 14, Aunuu, Tutuila, Samoa. BPBM 83256. a, pallial region, b, genitalia. c, interior of penis with muscular collar not slit open:
d-e, Sinployea clista, new species. Olomoana, Tutuila, Samoa. BPBM 84372. d, terminal genitalia. e, pallial region; f-h, Sinployea intermedia.
new species. West side of Swains Island. BPBM 186666. f, pallial region. #, genitalia. h, penis interior. Scale lines equal 1 mm. (MO).
tions it falls within the midrange of variation, height
1.35 mm., diameter 2.63 mm., H/D ratio 0.512, whorls
4Vs, D/U ratio 3.64. The spire is slightly more elevated
than normal, and this may be misidentified. If so, it
represents an undescribed species.
Description of soft parts. Pallial region (fig. 53a) about 3.3 mm.
long, rectal kidney lobe much longer than pericardia!, about 1.35
mm. long.
Genitalia (fig. 53b) without unusual features. Penis length
about 1.0 mm., internally (fig. 53c) penis with verge, collar, and
stimulatory pad. Latter drawing with collar shown intact, verge not
visible.
(Based on BPBM 83256, 1 example.)
Sinployea allecta (Cox, 1870).
Material from Ta'u, Upolu, and Savai'i is suffi-
ciently similar in size, shape, and sculpture (table
XXV) to be considered conspecific. The Swains Island
S. intermedia (fig. 52d-f) is conspicuously larger, has a
wider umbilicus, and averages two more apical spiral
cords. Sinployea aunuuana (fig. 52a-c) has the um-
bilicus much more widely open (mean D/U ratio 3.69 as
compared to more than 6.5 in allecta) and possesses
relatively conspicuous secondary spiral cording.
Sinployea complementaria (fig. 56a-f) is much, much
larger (table XXIV) and has more widely spaced radial
ribs.
Taxonomic treatment of the S. allecta material is
partly arbitrary. The populations on Ta'u are subspeci-
fically distinct, differing significantly in having both a
higher spire and a wider umbilicus. Usually a lower
spire means a wider umbilicus, and a higher spire ac-
companies a narrower umbilicus. Hence, the shifts in
the Ta'u populations are separate rather than being
linked. Subspecific status has been chosen because the
similarities are many. Populations on Savai'i and
Upolu show no significant differences in size or shape,
but do have markedly distinctive rib counts and spac-
ing (table XXV). The Upolu material comes from the
1860-1870 period, whereas the Savai'i material was
collected in 1965. As discussed below, only subadult S.
allecta were taken on Upolu in 1965. The significance
of this sculpture variation remains uncertain. In treat-
SYSTEMATIC REVIEW
125
ing the Savai'i and Upolu populations as belonging to
one subspecies, I am deferring judgment as to their
exact relationship. New collections of adults from
Upolu are needed to settle this problem.
Sinployea allecta allecta (Cox, 1870). Figures 54a-c,
55a-e.
Helix allecta Cox, 1870, Proc. Zool. Soc. London, 1870, p. 81 found
on the mountains under decayed wood, Upolu, Samoa; Pfeiffer,
1876, Monog. helic. viv., 7, p. 162.
Champa allecta (Cox), Tryon, 1886, Man. Conchol., (2) 2, p. 210
name only.
Patula allecta (Cox), Garrett, 1887, Proc. Acad. Nat. Sci., Philadel-
phia, 1887, p. 130.
Diagnosis. Shell larger than average, diameter 2.63-3.62 mm.
(mean 3.13 mm.), with 3%-4'/2 rather tightly coiled whorls. Apex and
early spire usually flat, sometimes slightly and evenly elevated,
lower spire descending slightly, body whorl descending moderately,
spire protrusion about VK body whorl width, H/D ratio 0.511-0.622
(mean 0.571). Apical sculpture of 7- 10 (mean 8.60) rather fine spiral
cords. Postnuclear whorls with narrow, rounded, rather prominent,
and protractively sinuated radial ribs that are variable in number
and spacing, 56121 (mean 89.1) on the body whorl, whose in-
terstices are 2-5 times their width. Ribs/mm. 5.83-11.98 (mean
9.00). Microsculpture of exceedingly fine radial riblets, 4-12 between
each pair of major ribs, crossed by slightly finer and more crowded
spiral riblets. No secondary spiral cording visible at 96 x magnifica-
tion. Umbilicus narrow, U-shaped, last whorl decoiling more rapidly,
slightly constricted by columellar lip expansion, contained 5.93-9.80
times (mean 7.85) in the diameter, margins rounded. Whorls almost
evenly rounded on outer margins, slightly flattened laterally above
periphery, aperture inclined about 15 from shell axis.
The narrower umbilicus and lower spire protru-
sion readily separate Sinployea allecta allecta (fig.
54a-c) from S. a. tauensis (fig. 54d-f). Sinployea inter-
media (fig. 52d-f) is larger and has a higher spire and
much wider umbilicus. Other Samoan species differ
radically in size and umbilical proportions. Of ex-
tralimital species with narrow umbilici, the Lau Ar-
chipelago S. angularis (fig. 64d-f) and S. recursa (fig.
64a-c) both have reduced radial ribbing; the Tahitian
S. montana (fig. 41a-c) is much smaller.
Description. Shell smaller than average with 3% loosely coiled
whorls. Apex slightly protruding, spire flat, last whorl descending a
little, H/D ratio 0.554. Body whorl quite obese, sutures moderately
deeply impressed, whorls strongly rounded above and on outer mar-
gins. Embryonic whorls 1%, sculpture of 9 relatively inconspicuous
spiral ribs. Postnuclear whorls with moderately prominent, rounded,
closely spaced, protractively sinuated radial ribs, 86 on the body
whorl, whose interstices are usually slightly less than twice their
width. Microsculpture of 4-8 very fine radial riblets between each
pair of major ribs, and even finer and more crowded spiral riblets.
Body whorl evenly rounded on outer margin with slightly protruded
basal lip coming to the evenly rounded umbilical margin. Umbilicus
partially constricted by inward expansion of columellar lip, narrowly
U-shaped, regularly decoiling, contained 5.93 times in the diameter.
Color yellowish white with relatively numerous reddish flammula-
tions. Aperture circular with protruding basal margin, inclined
about 15 from shell axis. Height of lectotype 1.53 mm., diameter
2.73 mm.
Lectotype. Samoa: Upolu. Collected on the moun-
tains under decayed wood. AMS C. 63457.
Range. Upolu and Savai'i, Western Samoa.
Paratypes.AMS C. 63458.
Material. Upolu (60 specimens, BPBM 106246,
BPBM 115349, BPBM 167430, AMS C.34756, AMS
C.63457-8, Zurich, FMNH 117267, FMNH 160001,
SMF 165706, NMWC): Togitogiga forest area (Station
10) at 50 ft. elevation (1 specimen, FMNH 153062); foot
of southeast peak (Station 7), Mt. Tafua Upolu at 1,200
ft. elevation (3 specimens, FMNH 153082); gully to foot
of Mt. Siga'ele (Station 23) at 2,300 ft. elevation (1
specimen, FMNH 153426).
Savai'i: heavy forest about 5 miles (Station 30)
northwest of Vailoa at 600 ft. elevation in heavy forest
(20 specimens, FMNH 153039); about Vz mile inland, 5
miles east of Vailoa (Station 38) on road to Salelologa
wharf in open bush (66 specimens, FMNH 153371,
FMNH 153091).
Remarks. The type specimens are all subadult,
and Cox's whorl count of 4'/2-5 is in error. In older
collections, specimens of this species are usually
labeled as S. complementaria. Numerous specimens
were collected by Graeffe in the early 1870s and are
rather widely distributed in museum collections. This
was after Mousson had published on Samoan material.
Hence, they were not mentioned in the previous litera-
ture.
Although the material in historical collections and
the specimens obtained by Solem and Price in 1965
agree almost exactly in dimensions and shape (table
XXV), there is a marked dichotomy in ribbing. The
specimens collected by Cox and Graeffe in the 1860s
and 1870s on Upolu have a mean of 99.3 ribs on the
body whorl and 9.90 ribs/mm. Those from Savai'i (ta-
bles XXIV- XXV) average only 75.9 ribs on the body
whorl and 7.84 ribs/mm. Unfortunately, only subadult
material was obtained on Upolu in 1965. The specimen
from Togitogiga (Station 10) has the typical Upolu rib-
bing, but those from Mt. Tafua Upolu (Station 7) and
near Mt. Siga'ele (Station 23) have the more widely
spaced ribbing found in the Savai'i examples.
Description of the soft parts. Foot and tail longer than shell
diameter, bluntly rounded anteriorly, slightly tapering posteriorly.
Sole undivided, pedal grooves prominent, suprapedal larger; no
caudal horn or middorsal groove present. Head projecting slightly in
front of foot. Ommatophores typical. Gonopore in normal position.
Body color yellow-white, with a grayish suffusion on neck, head,
and back of tail.
Mantle collar (fig. 55d) without glandular extension onto pallial
roof, edge swollen and providing a flap to narrow pneumostome.
Anus (A) and external ureteric pore (KX) typical.
Pallial region extending % whorl apically. Lung roof clear, with-
out granulations. Kidney (K) with lobes subequal, rectal lobe 1.45
mm. long, clearly overlapping hindgut. Base slightly indented, inner
side dished by organs above pallial cavity. Ureter (KD) reflexed with
very little roof space between arms (fig. 55a-b), narrow, widening
slightly along hindgut. Heart (H) more than half length of kidney,
angled slightly from hindgut. Principal pulmonary vein (HV) simple,
unbranched, angling toward pneumostome. Hindgut (HG) arising
just above kidney, not tapering.
Ovotestis with one or two clumps of palmately clavate acini
lying parallel to whorl sides. Hermaphroditic duct (fig. 55c, GD) nar-
row at first, expanded and iridescent, sharply reflexed and narrowed
at apex of albumen gland, following surface of gland to base of talon
head, entering latter basally. Albumen gland (GG) elongately ovate,
surface indented by intestinal loops and spermathecal head. Talon
(GT) with ovately globular, iridescent head partly buried in albumen
FIG. 54. a-c, Sinployea allecta allecta (Cox), Upolu, Samoa. BPBM 167430; d-e, Sinployea allecta tauensis, new subspecies. Station 188,
Faleasao, Ta'u, Samoa. Holotype. BPBM 187753. Scale line equals 1 mm. (MM).
126
HV
SYSTEMATIC REVIEW
127
PP
FIG. 55. Anatomy of Sinployea allecta allecta (Cox). Station 38,
Vailoa, Savai'i, Samoa. FMNH 153091. a, external view of pallial
region; b, internal view of pallial region; c, genitalia; d, mantle
collar at parietal-palatal angle showing lobe development; e, interior
of penis. Scale lines equal 1 mm. (MM).
gland, shaft very slender, joined by hermaphroditic duct just below
head expansion. Prostate (DG) of few large acini opening into a
groove on upper uterine chamber wall. Uterus (UT) bipartite, upper
chamber very thin-walled, lower with thick, glandular surface.
Vas deferens (VD) reflexing up from penioviducal angle, enter-
ing epiphallus (E) through a "valve" pilaster (fig. 55e). Epiphallus
more than half the length of penis, entering penis apically. Penial
retractor (PR) arising on diaphragm, inserting in "U" around
epiphallus-penis junction on side opposite epiphallic ascension. Penis
(P) 1.3-1.5 mm. long, not tapering basally. Internally (fig. 55e) with
vergic papilla enlarged and elongated, fastened to muscular collar on
one side, opening to epiphallus occupying small portion of length.
Main stimulatory pad lower and less sharply outlined than in most
species. Atrium (Y) narrow and relatively short, without clear pilas-
ter arrangement.
Free oviduct (UV) with thick muscular walls, internally with
very narrow channel. Spermatheca (S) with expanded head lying
next to albumen gland, expanded basally with same pilaster ar-
rangement found in S. complernentaria. Vagina (V) as in S. com-
plementaria.
Free muscle system without unusual features.
Buccal mass elongately oval, small generative sac. Buccal re-
tractors not split, inserting on lower posterior margin of buccal mass.
Esophagus a narrow tube extending just apicad of pallial cavity.
Stomach extending % whorl apically, anterior expansion very sud-
den. Intestinal looping normal, compressed into less than V whorl.
Salivary glands uniting posteriorly above esophagus.
(Based on FMNH 153091, 5 adult, expanded examples, 3.16-
3.29 mm. in diameter, with 4'/8-4% whorls.)
Sinployea allecta tauensis, new subspecies. Figure
54d-e.
Diagnosis. Shell slightly larger than average, diameter 2.70-
3.39 mm. (mean 3.00 mm.), with 3%-4'/ somewhat tightly coiled
whorls. Apex and spire noticeably and evenly elevated, body whorl
descending much more rapidly, spire protrusion more than '& body
whorl width, H/D ratio 0.541-0.616 (mean 0.577). Apical whorls
worn in all material, traces of fine spiral cords visible in most. Post-
nuclear whorls with fine, low, rounded, protractively sinuated radial
ribs, 61-101 (mean 79.8) on the body whorl, whose interstices are
2-4 times their width. Ribs/mm. 6.20-10.51 (mean 8.31). Micro-
sculpture of extremely fine radial riblets, 6-12 between each pair of
major ribs, crossed by even finer and more crowded spiral riblets. No
secondary spiral cording. Umbilicus narrow, V-shaped, last whorl
decoiling more rapidly, contained 4.72-9.55 times (mean 6.80) in the
diameter, margins rounded. Whorls almost evenly rounded, not or
only slightly flattened laterally above periphery.
The nominate race of Sinployea allecta (fig. 54b-c)
has a narrower umbilicus and slightly lower spire than
S. a. tauensis (fig. 54e-f). The only Polynesian species
apt to be confused with this is the Cook Island S. av-
anaensis (fig. 45d-f), which is much less elevated and
has a wider umbilicus.
Description. Shell rather large, with slightly more than 4
rather tightly coiled whorls. Apex and spire slightly and evenly ele-
vated, last whorl descending more rapidly, H/D ratio 0.545. Em-
bryonic whorls 1%, sculpture of fine spiral cords visible in suture,
eroded over most of surface. Postnuclear whorls with prominent,
rounded, variably spaced, protractively sinuated radial ribs, 76 on
the body whorl, whose interstices are 2-4 times their width, and
which become very irregular near aperture. Microsculpture of ex-
tremely fine radial riblets, 5-12 between each pair of major ribs,
crossed by slightly finer and more crowded spiral riblets. No secon-
dary spiral cording visible at 96 x magnification. Sutures deep,
whorls evenly rounded on outer margins. Umbilicus narrow, V-
shaped, last whorl decoiling more rapidly, contained 5.94 times in
the diameter, slightly narrowed by columellar lip expansion. Color
reddish horn on spire, last whorl separating into basal zones and
128
SOLEM: ENDODONTOID LAND SNAILS
broad, irregular, sinuated reddish flammulations that coalesce on
shell base. Aperture almost circular, with evenly rounded outer
margins, inclined about 20 from shell axis. Height of holotype 1.81
mm., diameter 3.32 mm.
Holotype. Samoa: Ta'u, Station 188, Faleasao,
700 ft. inland at 12 ft. elevation. Collected by Wray
Harris on September 24, 1937. BPBM 187753.
Range. Ta'u, Manu'a Group, American Samoa.
Paratypes. Ta'u: Talavagi ridge and summit
(Station 75) from I fi2-Vs mile inland at 450-500 ft. ele-
vation (4 specimens, BPBM 171106, BPBM 181998,
BPBM 182095); Siufaga (Stations 76, 184), 250-600 ft.
inland at 400-600 ft. elevation (12 specimens, BPBM
171073, BPBM 182055, BPBM 182400, BPBM 182410,
BPBM 182444, BPBM 187635, BPBM 187640, BPBM
187653); Maia (Station 87), l /iz mile inland at 75 ft.
elevation (1 specimen, BPBM 182364); Faleasao (Sta-
tions 188, 189), 600-700 ft. inland at 12 ft. elevation (8
specimens, BPBM 187766, BPBM 187781, BPBM
187788, BPBM 188357, ex BPBM 187736, ex BPBM
188277).
Remarks. All known material consists of dead
specimens sorted from rubble sweepings. Normally a
higher spire and narrower umbilicus are correlated
changes, so that the more protruded spire and wider
umbilicus of Sinployea allecta tauensis, compared with
the nominate race, are doubly significant. Rib spacing
and counts in S. a. tauensis are very similar to those
found in the Savai'i population and rather different
from the Upolu examples (table XXVI).
TABLE XXVI. - LOCAL VARIATION IN SAMOAN SINPLOYEA, II.
NUMBER OF
SPECIMENS
allecta tauensis
RIBS
77.34.18
(66-95)
HEIGHT
1.760.048
(1.61-1.97)
DIAMETER
3.050.076
(2.76-3.32)
H/D RATIO
0.577+0.0096
(0.542-0.606)
Faleasao
Siufa
ga
4
88.77.89
(74-101)
1.72+0
(1
.059
.61-1.88)
3.040
(2
.141
.70-3.39)
0.5660
(0
.0120
.543-0
.598)
intermedia
BPBM
186666 14
81.74.93
(65-96)
2.030
(1
.032
.82-2.19)
3.490
(3
.042
.21-3.81)
0.581+0
(0
.0062
.534-0
.616)
complement aria
Sta.
FMNH
39, 2000' 8
153386, -419
66.3+1.67
(59-73)
2.330
(2
,062
,11-2.57)
4.71+0
(4
.150
.31-5.20)
0.497+0
(0
.0113
.458-0
,543)
Sta.
FMNH
-168
18, 600-800' 37
153016, -24,
,-176, -557
69.9tl.24
(54-87)
2.430
(2
.025
11-2.70)
4.81+0
(4,
.038
,34-5.23)
0.5050
(0
.0040
.464-0
.563)
Sta.
FMNH
26, 2450-2500' 8
153047, -99
62.9+2.40
(58-78)
2.73+0.070
(2.53-3.03)
5.110.
(4
,124
.57-5.53)
0.534+.0,
(0,
,0117
473-0,
561)
Sta.
FMNH
24, 2300' 3
153619
53.01.00
(52-55)
2.820.
(2.
105
63-2.99)
5.210,
(5,
090
03-5.33)
0.54110,
(0,
0110
523-0.
562)
Sta. 13
FMNH
,750' 5
153380, -408
79.8+2.98
(73-88)
2.83+0.
(2.
119
47-3.09)
5.32+0,
(4,
217
57-5.92)
0.532+0.
(0.
0124
506-0.
577)
WHORLS
all.tau
Faleasao 4+(3 7/8-4 1/4)
Siufaga
4-(3 3/4-4 1/8)
TABLE XXVI, CONTINUED
UMBILICUS
0.49+0.047
(0.36-0.69)
0.44+0.049
(0.33-0.56)
D/U RATIO
6.45+0.515
(5.53-8.45)
7.140.483
(6.06-8.20)
APICAL CORDS
BPBM
compleme .
Sta. 39
Sta. 26
Sta. 24
Sta. 13
4 3/8+(4 1/4-4 3/4) 0.64+0.019
(0.53-0.79)
4 5/8-(4 1/2-4 7/8) 0.93+0.046
(0.76-1.18)
Sta. 18 4 5/8+(4 3/8-5)
0.99+0.016
(0.82-1.15)
4 7/8-(4 1/2-5 1/4) 1.01+0.052
(0.82-1.22)
5-(4 3/4-5)
4 7/8-(4 5/8-5)
0.99
1.070.662
(0.92-1.18)
5.43+0.123 10.10.31
(4.59-6.12) (8-12)
5.13+0.149
(4.61-6.00)
4.890.060 9.82+0.48
(4.18-5.77) (8-13)
5.14+0.202 13
(4.22-6.08)
5.28+0.160
(5.10-5.40)
4.9710.109
(4.61-5.29)
SYSTEMATIC REVIEW
129
Sinployea complementaria (Mousson, 1865). Fig-
ures 56a-f, 57a-f.
Patula complementaria Mousson, 1865, J. de Conchyl., 13, pp.
168-169, 431, pi. 14, fig. 5 Upolu, Samoa; Mousson, 1869, J. de
Conchyl., 17, p. 333; Garrett, 1887, Proc. Acad. Nat. Sci.,
Philadelphia, 1887, pp. 129-130 behind Apia, Upolu, Samoa.
Helix complementaria (Mousson), Pfeiffer, 1868, Monog. helic. viv.,
5, p. 157; Pfeiffer, 1876, Monog. helic. viv., 7, p. 164.
Pitys complementaria (Mousson), Pease, 1871, Proc. Zool. Soc. Lon-
don, 1871, p. 474.
Helix (Patula) complementaria (Mousson), Tryon, 1887, Man.
Conchol., (2) 3, pp. 40-41, pi. 8, fig. 57.
Endodonta (Champa) complementaria (Mousson), Pilsbry, 1894,
Man. Conchol., (2) 9, p. 35.
Diagnosis. Shell extremely large, diameter 4.31-5.92 mm.
(mean 4.92 mm.), with 4% 5!4 normally coiled whorls. Apex and
early spire barely to strongly elevated, last whorl usually descending
much more rapidly, spire protrusion quite variable, usually about l h
body whorl width, H/D ratio 0.458-0.648 (mean 0.517). Apical
sculpture of 8-13 (mean 10.08) fine spiral cords, frequently eroded.
Postnuclear whorls with high, narrow, prominent, strongly protrac-
tively sinuated radial ribs, 52-95 (mean 69.1) on the body whorl,
whose interstices are 3-7 times their width. Ribs/mm. 3.11-5.79
(mean 4.50). Microsculpture of extremely fine radial riblets, 9 to
more than 20 between each pair of major ribs, crossed by slightly
finer and more crowded spiral riblets. No secondary spiral cording.
Umbilicus open, V-shaped, last whorl decoiling more rapidly, con-
tained 4.18-6.08 times (mean 4.99) in the diameter, margins
rounded. Whorl contour slightly more rounded than average, flat-
tened laterally above periphery, slightly on basal margin, aperture
inclined about 20 from shell axis.
The extremely large size, comparatively few radial
ribs, and a high number of microradial riblets effec-
tively separate even juvenile examples of Sinployea
complementaria (fig. 56a-f) from the other Samoan
species. Of the larger Rarotongan species, only S.
tenuicostata (fig. 50d-f), which has a flat spire, fewer
and larger microradial riblets plus never more than 4'/2
whorls, might be confused. The Tongan Tuimalila (fig.
78a-f) are even larger, but the looser whorl coiling and
reduced apical sculpture of that genus readily sepa-
rates them.
Description. Shell very large, with slightly more than 4V4 nor-
mally coiled whorls. Apex barely emergent, whorls of spire descend-
ing rapidly, H/D ratio 0.648. Apical whorls l'/2, sculpture eroded.
Postnuclear whorls with narrow, moderately widely spaced, protrac-
tively sinuated radial ribs, 73 on the body whorl, whose interstices
are 3-5 times their width. Microsculpture a lattice of very fine and
crowded radial riblets crossed by distinctly finer and slightly more
crowded spiral riblets. No secondary spiral cording. Sutures moder-
ately impressed, whorls slightly shouldered above, strongly flattened
laterally above periphery and on basal margin. Umbilicus V-shaped,
narrow, partially closed by reflection of basal lip, contained 6.00
times in the diameter. Color light yellow-brown with irregular red-
dish flammulations, generally parallel to the line of growth. Aper-
ture subcircular, flattened laterally above periphery and on basal
margin, inclined about 15 from the shell axis. Edge of lip broken.
Height of lectotype 3.32 mm., diameter 5.14 mm.
Lectotype. Samoa: Upolu. Collected by Graeffe.
Zoologisches Museum der Universitat Zurich.
Range. Upolu, Western Samoa.
Paratypes. Zurich.
Material. Samoa (2 specimens, BPBM 161):
Upolu (14 specimens, Zurich, IRB); foot of Mt. Solaua
(Station 18) at 600-800 ft. elevation on stilts ofFicus
or under fallen bark and rotting wood (59 specimens,
FMNH 153016, FMNH 153024, FMNH 153165,
FMNH 153168, FMNH 153176, FMNH 153557);
Fagaloa Pass (Station 13) at 750 ft. elevation along
ridge crest under rotting wood (6 specimens, FMNH
153380, FMNH 153408); Le Mafa Pass at 900 ft. eleva-
tion under birdnest fern (2 specimens, DMW M-4002,
collected by R. A. Cumber on February 10, 1955); top of
range (Station 39), Mt. Solaua at 2,000 ft. elevation in
heavy forest (18 specimens, FMNH 153386, FMNH
153419); Afiamalu, 6 miles inland at 2,100 ft. elevation
(3 specimens, BPBM 186029, BPBM 186214); % mile
above Afiamalu seismographic station (Stations 2, 8)
in secondary forest (2 specimens, FMNH 153050,
FMNH 153089); south slope, Mt. Siga'ele (Station 24)
at 2,300 ft. elevation in heavy forest (3 specimens,
FMNH 153619); Afiamalu-Lake Lanuto'o track (Sta-
tion 26) at 2,450-2,500 ft. elevation under rotting wood
in heavy forest (9 specimens, FMNH 153047, FMNH
153099); rim of Lake Lanuto'o crater (Station 19) at
2,500 ft. elevation under dead leaves and rotting wood
(18 juvenile specimens, FMNH 153133, FMNH
153424).
Remarks. The lectotype is the highest specimen
seen, but closely matches the type figures and has the
characteristic sculpture well preserved. Spire elevation
in the juvenile example (fig. 56e) is slightly less than
average.
Samples collected in 1965 showed rather consider-
able variation, some of which may be environmentally
correlated. The very large shells (table XXVI) from
Fagaloa Pass come from an area of very high rainfall
that also is continually swept by a moist sea breeze.
Hence humidity would remain continually high, and
conditions for growth would be exceptionally favorable.
Specimens from Mt. Siga'ele (Station 24) were taken
under litter in heavy fog forest. Their low rib count and
wide rib spacing may reflect the constancy of this
niche. Only juvenile examples were taken on the rim of
Lanuto'o crater (Station 19). Almost all examples from
Station 18 at the foot of Mt. Solaua were collected
around the base of a single huge Ficus that was iso-
lated from the forested slopes by a partly overgrown
banana patch. Specimens of Sinployea complementaria
were most frequent under starting bark on stilt roots
from 2-5 ft. above ground level. A number were taken
from under accumulated debris in the center of this
tree, and only a few under fallen branches lying in
grass beneath the canopy spread. None were found in
the banana patch or lower forest reaches. Collecting
1,200 ft. higher on top of Mt. Solaua (Station 39) pro-
duced specimens essentially identical in measure-
ments (table XXVI) and ribbing.
Without much more material, the question of
geographic variation cannot be settled. The size
difference between the Mt. Solaua and Fagaloa or
Siga'ele specimens is large and significant, but there is
no obvious geographic pattern. The occurrence of S.
complementaria is strictly colonial, the specimens from
Mt. Solaua, Fagaloa, and the Lake Lanuto'o areas hav-
ing been obtained in a fraction of the collecting time
devoted to the Afiamalu area with its two examples.
Sinployea complementaria was found in both pro-
a
FIG. 56. Sinployea cornplementaria (Mousson). a-c, Upolu, Samoa. Lectotype of Patula complementaria Mousson, 1865. Zoologisches
Museum Universitat Zurich; d-f, Afiamalu, 2,100 ft., Upolu, Samoa. Juvenile. BPBM 186214. Scale lines equal 1 mm. (SG).
130
SYSTEMATIC REVIEW
131
foundly disturbed (Fagaloa and foot of Mt. Solaua) and
undisturbed vegetation zones.
Description of soft parts. Foot and tail equal in length to shell
diameter, slightly tapering posteriorly, truncated anteriorly. Sole
undivided, pedal grooves high on foot, suprapedal weaker, no caudal
horn or middorsal groove. Slime network typical. Gonopore a long,
vertical slit behind right rhinophore and below right ommatophore.
Body color yellow-white, with light gray speckling on head and
neck, ommatophores not darker.
Mantle collar (MC) swollen and rolled, no glandular extension
onto pallial roof, a dark gray streak on expanded portion. Pneumo-
stome masked by expansion of collar, no mantle lobe development.
Anus (A) and external ureteric pore (KX) opening beside each other
just inside pneumostome.
Pallial region (fig. 57a) extending % whorl apically. Lung roof
with black specklings along edges of pulmonary vein, otherwise
clear. Kidney (K) bilobed, 2.57 mm. long, rectal arm slightly longer
than pericardial and crossing hindgut onto parietal wall. Ureter
(KD) typical, a narrow strip of lung roof visible between arms. Heart
(H) slightly more than Vs length of kidney, angled to hindgut. Princi-
pal pulmonary vein (HV) unbranched, fading out just short of mantle
collar. Hindgut (HG) deflecting from parietal-palatal margin just
before end of kidney.
Ovotestis (fig. 57b, G) a single clump of palmately clavate al-
veoli, lying parallel to whorl sides, occupying Vs whorl above stomach
apex. Hermaphroditic duct (GD) slender at first, expanded and
iridescent in middle, markedly kinked just before albumen gland,
embedded and narrow before talon. Albumen gland (GG) with rather
large acini, surface indented by intestinal loops and spermathecal
head. Talon (fig. 57d, GT) with globular head, hermaphroditic duct
inserting at base. Carrefour imbedded, a slightly expanded tube from
neck of talon, opening into prostate-uterus. Prostate (DG) of large
acini opening into groove on side of uterus, acini complexly folded.
Uterus (UT) bipartite, upper chamber with very thin walls.
Vas deferens (VD) a broad tube with heavily glandular walls to
penioviducal angle, where it narrows abruptly to ascend along side of
penis. Epiphallus (E) large, twisted, entering head of penis laterally,
with a peculiar Y-shaped valve (fig. 57e) at entrance of vas deferens,
lower portion with glandular pilasters, entering penis laterally on
head (fig. 57c). Penial retractor (PR) short, thick, arising on dia-
phragm, inserting onto head of penis. Penis (P) about 1.75-1.85 mm.
long, club-shaped, gradually tapering to atrium, internally (fig. 57c)
with large verge (PV) through which epiphallic pore (EP) enters
medially, a reduced sphincter pilaster and a moderately large, typi-
cal pocket pilaster. Atrium (Y) of average length, with weak lon-
gitudinal pilasters.
Free oviduct (UV) a long, thick muscular tube with an apical
muscle attaching to right tentactular retractor, internally (fig. 57f)
with heavy glandular walls ending in a hemispherical protrusion
with subcentral pore (UVO) just at junction with spermathecal base,
externally smoothly muscled. Spermatheca (S) with head lying next
to albumen gland, slender shaft bound to diaphragm tissue to base of
uterus, then grossly expanded to vagina level, with complex pilasters
forming a circular ring at area of vagina, with a flap obviously capa-
ble of closing the spermathecal opening (SO). Vagina (V) quite
thin-walled, with weak internal pilasters.
Free muscle system typical, muscle from free oviduct quite
prominent. Right ommatophoral retractor passing through peni-
oviducal angle.
Buccal mass and esophagus without unusual features. Stomach
occupying a little more than % whorl, shape and expansion typical.
Intestinal looping occupying less than Vs whorl above pallial cavity
apex.
Salivary glands fused across top of esophagus.
Central tooth of radula distinctly smaller than 1st lateral,
tricuspid with mesocone extending anterior to basal plate edge. Lat-
erals 4-5, with endocone increasing regularly, transitions to margi-
nals marked by shift in support size and increasing size of endocones.
Marginals 6-7, outer ones with endocone and mesocone subequal,
ectocone always much smaller, no evidence of ectoconal splitting.
(Based on FMNH 153165, 4 whole individuals, 4.9-5.1 mm. in
diameter.)
Sinployea intermedia, new species. Figures 52d-f,
53f-h.
Diagnosis. Shell large, diameter 3.21-3.81 mm. (mean 3.49
mm.), with 4'/4-4% rather tightly coiled whorls. Apex and spire mod-
erately and evenly elevated, last whorl descending more rapidly,
spire protrusion about '& body whorl width, H/D ratio 0.534-0.616
(mean 0.581). Apical sculpture of 9-13 (mean 10.57) very fine spiral
cords. Postnuclear whorls with low, rounded, prominent, protrac-
tively sinuated radial ribs, 63-96 (mean 82.1) on the body whorl,
whose interstices are usually 3-5 times their width. Ribs/mm.
5.75-9.16 (mean 7.66). Microsculpture of fine radial riblets, 5-10
between each pair of major ribs, crossed by slightly finer and more
crowded spiral riblets. No secondary spiral cording. Umbilicus nar-
row, U-shaped, last whorl decoiling more rapidly, contained 4.59-
6.12 times (mean 5.43) in the diameter, margins rounded. Whorls
only slightly flattened laterally above periphery, strongly rounded
on basal margin.
Sinployea intermedia (fig. 52d-f) has a distinctly
more protruded spire, wider umbilicus, slightly greater
number of apical cords and is larger than any of the S.
allecta (fig. 54a-e) races. Society and Cook Island
species of the same size are more widely umbilicated
and have quite different ribbing structure.
Description. Shell larger than average, with slightly more
than 4'/2 moderately tightly coiled whorls. Apex and spire moder-
ately and evenly elevated, body whorl descending more rapidly, H/D
ratio 0.592. Embryonic whorls l'/2, sculpture of 8 relatively weak
spiral ribs with irregularly radially ribbed last quarter whorl. Re-
maining whorls with low, rounded, moderately widely spaced, pro-
tractively sinuated radial ribs, 90 on pregerontic portion of body
whorl, whose interstices are about 2-3 times their width. Micro-
sculpture of extremely fine microradial riblets, 5- 10 between each
pair of major ribs, and smaller microspiral riblets forming a latticed
pattern. Sutures relatively deep, whorls strongly rounded above and
on basal margin, body whorl slightly deflected at aperture. Um-
bilicus narrowly U-shaped, becoming more widely open on last
whorl, contained 4.91 times in the diameter. Color light yellowish
white with numerous irregular reddish flammulations. Aperture cir-
cular, slightly compressed laterally above periphery by deflection of
lip, inclined about 15 from the shell axis. Height of holotype 2.11
mm., diameter 3.56 mm.
Holotype. Swains Island: 200 yd. inland on west
side at 20 ft. elevation. Collected under coconut husks
and fallen limbs by Elwood C. Zimmerman on August
21, 1940. BPBM 186666.
Range. Swains Island, American Samoa.
Paratypes. Swains Island: west side, 200-500 yd.
inland at 20 ft. elevation (45 specimens, BPBM
186666-8, BPBM 186679).
Remarks. Differences from Sinployea allecta are
adequately covered in the diagnosis above. Although
closely related to that species, I have no doubt that
Sinployea intermedia is distinct. Presence of an en-
demic species on Swains Island, a very isolated but
heavily vegetated atoll, was unexpected. Swains Island
is less than 20 ft. high and not over IVz mile in greatest
dimension (Bryan, 1942, p. 83).
132
SOLEM: ENDODONTOID LAND SNAILS
VD
UVO
FIG. 57. Anatomy of the Samoan Sinployea complementaria (Mousson). Station 18, Mt. Solaua, Upolu, Samoa. FMNH 153165: a, pallial
region; b, genitalia; c, interior of penis; d, detail of talon-carrefour area; e, junction of vas deferens and epiphallus; f, interior of spermatheca and
free oviduct junction. Scale lines equal 1 mm. (SH).
Description of soft parts. Foot and tail shorter than shell
diameter. Sole undivided, slightly tapering posteriorly. Pedal
grooves high on sides of foot, prominent.
Body color light yellow-white, no grayish markings.
Mantle collar (MC) thick and swollen in available material, no
glandular extension onto mantle roof.
Pallial region (fig. 530 extended length about '/a whorl. Lung
roof clear of granulations. Kidney (K) bilobed, rectal lobe longer than
pericardial, greatest length 1.65-2.00 mm., differing from S. allecta
in having less overlap of hindgut and greater lung roof visible be-
tween arms of ureter (KD). Heart (H) and principal pulmonary vein
(HV) typical. Hindgut (HG) following parietal-palatal margin for
moderate distance above kidney apex.
Ovotestis (fig. 53g, G) composed of one or two acinar clumps in
typical position. Hermaphroditic duct (GD), albumen gland (GG),
talon (GT), prostate (DG), and uterus (UT) without unusual features.
Vas deferens (VD) and epiphallus (E) as in Sinployea allecta.
Penial retractor (PR) as in S. allecta. Penis (P) 1.2-1.6 mm. long,
SYSTEMATIC REVIEW
133
tapering more on basal portion than inS. allecta, externally strongly
swollen on upper half. Internally (fig. 53h) with typical stimulatory
pad, vergic papilla as in S. allecta. Atrium (Y) very short, without
unusual features.
Free oviduct (UV), spermatheca (S), and vagina (V) typical; lat-
ter a trifle longer than in S. allecta.
(Based on BPBM 186666, several partial examples.)
WESTERN POLYNESIAN AND MICRONESIAN Sinployea
Whereas the Sinployea from Tonga, Hoorn Islands,
Ellice Islands, and Rotuma show many similarities, in-
clusion of the single species from Kusaie in the
Caroline Islands and a possibly introduced form from
Saipan in the Marianas is merely for convenience in
reducing the number of subheadings. Sinployea
kusaieana (fig. 63a-c) lacks gray coloration on the
head and ommatophores, has monochrome shell colora-
tion, a less protruded spire, fewer and more loosely
coiled whorls, and the umbilicus wider and V-shaped.
Two specimens collected on Saipan could represent an
introduced population, a mislabeled set, or another un-
described Micronesian species.
Of the remaining species, only Sinployea
pseudovicaria (fig. 61a-c) with its low H/D ratio and
wide umbilicus (table XXVII) is distinctive. The Tahi-
tian S. lamellicosta (Garrett) (fig. 40a-c) is virtually
identical in size and shape, but has a depressed spire,
averages less than half as many ribs, and has slightly
fewer whorls at a given diameter. Other widely umbili-
cated Sinployea are either much bigger (S. canalis, fig.
49a-c), have many more whorls (S. planospira, fig.
46d), or are much smaller (S. nissani, S. navutuensis).
Sinployea rotumana (E. A. Smith) (fig. 61d-f) perhaps
comes closest of any species to being median in its fea-
tures. Only the slightly more numerous and crowded
radial ribs depart from the median measurements out-
lined in Table XV. Sinployea ellicensis (fig. 62a-f) is a
rather small, normally elevated species with crowded
radial ribbing. Sinployea vicaria (Mousson) (fig. 58a-d)
is larger and with a rather elevated spire. Both of these
species have less marked lateral whorl flattening than
species from Eastern Polynesia. Although measure-
ments overlap, the round apertures and generally ele-
vated spires would allow immediate recognition in
mixed sets.
Because none of the species are sympatric, no for-
mal key has been prepared. The following geographic
notes will locate material of known taxa:
TABLE XXVII. - RANGE OF VARIATION IN WESTERN POLYNESIAN AND MICRONESIAN SINPLOYEA.
NAME
ellicensis
ellicensis
ellicensis
nukulaelaeana
vicaria
vicaria
(Mousson)
vicaria
paucicosta
rotumana
(Smith)
pseudovicaria
Vaitupu
Eua (?)
kusaieana
sp. (Saipan)
NUMBER OF
SPECIMENS
326
87
9
21
7
61
2
RIBS RIBS/MM. HEIGHT DIAMETER H/D RATIO
100.3(79-125) 13.23(10.55-16.69) 1.36(1.13-1.68) 2.47(2.20-2.81) 0.549(0.486-0.600)
84.5(72-102) 13.02(11.06-15.20) 1.11(1.06-1.16) 2.08(1.99-2.20) 0.533(0.496-0.561)
108.5(77-145) 12.65(8.72-15.91) 1.51(1.17-1.89) 2.73(2.14-3.15) 0.552(0.469-0.633)
77.1(51.98) 9.80(6.71-11.44) 1.45(1.18-1.78) 2.49(2.22-2.76) 0.585(0.500-0.659)
105.2(84-144) 11.74(9.79-14.15) 1.55(1.25-1.81) 2.84(2.60-3.24) 0.547(0.481-0.607)
108.1(82-127) 11.39(9.56-15.17) 1.57(1.41-1.74) 3.24(3.03-3.52) 0.486(0.454-0.525)
94.8(81-113) 8.93(8.43-9.77) 1.61(1.45-1.74) 3.37(3.06-3.68) 0.479(0.455-0.509)
91.1(84-103) 11.25(10.27-13.12) 1.32(1.19-1.49) 2.60(2.52-2.75) 0.506(0.462-0.542)
85.0(82-88) 11.44(11.20-11.67) 1.41(1.38-1.45) 2.37(2.34-2.40) 0.598(0.592-0.603)
APICAL
CORDS
SPIRE
ELEVATION
BODY WHORL
WIDTH
e.ell.
e.nuk.
v. pau.
rotum.
pseud.
Vait.
Eua
kusai.
sp.
WHORLS UMBILICUS D/U RATIO CORDS ELEVATION WIDTH SP/BWW
4 1/8(4-4 1/2) 0.56(0.43-0.76) 4.39(3.48-5.46) 9.42(8-12) 0.15(0.10-0.23) 0.82(0.72-0.92) 0.183(0.125-0.259)
3 7/8+(3 5/8-4) 0.48(0.44-0.59) 4.31(4.13-4.66) 11.67(11-13) 0.11(0.10-0.16) 0.68(0.64-0.72) 0.166(0.136-0.250)
4 1/4(3 3/4-4 3/4) 0.58(0.41-0.75) 4.47(3.88-5.80) 10.63(9-12) 0.21(0.10-0.38) 0.86(0.69-1.02) 0.251(0.111-0.417)
4 1/2(4 1/4-5) 0.69(0.49-0.72) 4.14(3.62-4.64) 11.50(10-12) 0.21(0.10-0.33) 0.79(0.69-0.89) 0.268(0.125-0.385)
4 1/4(4-4 5/8) 0.66(0.58-0.81) 4.32(3.93-4.86) 9.67(8-11) 0.17(0.07-0.25) 0.95(0.84-1.04) 0.175(0.073-0.268)
4 3/8+(4 1/4-4 5/8) 1.03(0.79-1.22) 3.17(2.89-4.05) 11.74(10-13) 0.10(0.07-0.13) 0.86(0.79-0.95) 0.121(0.083-0.160)
4 1/2(4 1/4-4 3/4) 1.08(0.99-1.25) 3.13(2.95-3.30)
3 7/8+(3 3/4-4 1/8) 0.74(0.69-0.79) 3.51(3.21-3.67) 9.20(8-10) 0.12(0.10-0.16) 0.79(0.72-0.86) 0.150(0.125-0.208)
3 7/8+ 0.53(0.49-0.56) 4.52(4.30-4.73)
134
SOLEM: ENDODONTOID LAND SNAILS
Tonga and Hoorn Islands: S. vicaria (Mousson,
1871)
Sinployea vicaria paucicosta, new subspecies
on southern Vava'u
Rotuma Island: S. rotumana (E. A. Smith, 1897)
Ellice Islands
Vaitupu: S. pseudovicaria, new species
(widely umbilicated)
Funafuti: S. ellicensis ellicensis, new sub-
species
Nukulaelae: S. ellicensis nukulaelaeana, new
subspecies
Caroline Islands
Kusaie: S. kusaieana, new species
Mariana Islands
Saipan: Sinployea sp.
Sinployea vicaria (Mousson, 1871).
Several small samples from scattered islands of
the Tonga and Hoorn group collected by Graeffe in the
1860s and 1870s, plus material from Vava'u, Ton-
gatapu, and Eua collected by L. Price in 1966 agree
closely in proportions (table XXVIII). Four samples
from Vava'u have a distinctive pattern of more widely
spaced ribbing and are separated as a subspecies,
Sinployea vicaria paucicosta (fig. 58d). Size differences
among the remaining samples may be attributed to
sample bias and/or ecophenotypic dwarfing (p. 108).
They are grouped as a single subspecies. It is quite
possible that more ample material, particularly from
Futuna, will cause a revision in this judgment.
General appearance and sculpture are much more
similar to the smaller Cook Island Sinployea, S. atien-
sis, S. andrewi, and S. peasei, than to any of the Sa-
moan or Fijian species.
Sinployea vicaria vicaria (Mousson, 1871). Figures
58a-c, 60a-d.
Patula (Patula) vicaria Mousson, 1871, J. de Conchyl., 19, pp.
11-12, pi. 3, fig. 2 Futuna, Hoorn Group; Hufena on Hapai
(probably is Uoleva Id., Haapai Group) and Vavao, Tonga Is-
lands.
Helix vicaria (Mousson), Pfeiffer, 1876, Monog. helic. viv., 7, p. 187.
Helix modicella (Ferussac), Tryon, 1887, Man. Conchol., (2) 3, p.
38, pi. 8, figs. 31-32.
Rhytida (Ouagapia) vicaria (Mousson), Mollendorff, 1903, Syst.
Conchyl. Cab., I, 12b, pp. 83-84, pi. 13, figs. 16-17.
Diagnosis. Shell variable in size, generally slightly smaller
than average, diameter 2.14-3.15 mm. (mean 2.73 mm.), with
3%-4% rather tightly coiled whorls. Apex and spire markedly and
almost evenly elevated, last whorl descending moderately to much
more rapidly, spire protrusion averaging V* body whorl width, H/D
ratio 0.469-0.633 (mean 0.552). Apical sculpture of 9-12 (mean
10.63) very fine spiral cords. Postnuclear whorls with narrow,
rounded, crowded, rather strongly protractively sinuated radial ribs,
77-145 (mean 108.5) on the body whorl, whose interstices are 1-3
times their width. Ribs/mm. 8.72-15.91 (mean 12.65). Micro-
sculpture of very fine radial riblets, 5-8 between each pair of major
ribs, barely visible microspiral riblets, and rather closely spaced sec-
ondary spiral cords equal in size to microradial riblets. Umbilicus
rather narrow, U-shaped, last whorl decoiling more rapidly, con-
tained 3.88-5.80 times (mean 4.47) in the diameter, margins
rounded. Whorls less strongly flattened laterally than in Society or
Cook Island species.
Sinployea vicaria (table XXVII) is very similar to
S. atiensis (table XIX) in all measurements, differing
only slightly in ribbing and umbilical coiling. In S.
atiensis (fig. 44d-f) the umbilicus decoils regularly, the
secondary spiral cording is noticeably wider spaced,
and the whorl contour is more strongly flattened later-
ally. Sinployea vicaria (fig. 58a-c) usually has a more
rapid decoiling of the umbilicus and slightly more
crowded ribbing. Sinployea pseudovicaria (fig. 61a-c)
is immediately separable by its nearly flat spire and
much more widely open umbilicus. Sinployea rotu-
mana (fig. 61d-f) has the body whorl strongly flattened
laterally and more crowded secondary spiral cording.
Sinployea neglecta (fig. 41d-f) has finer ribs, fewer
microradial riblets, a V-shaped umbilicus, and greater
lateral whorl flattening.
Description. Shell of average size, with 4V* normally coiled
whorls, apex barely emergent, whorls of spire descending moder-
ately, last whorl more rapidly, H/D ratio 0.538. Apical whorls 1%,
sculpture of 11 fine, moderately widely spaced spiral ribs with a faint
microsculpture of radial ribs and a secondary intrusion of radial
growth wrinkles near the end. Postnuclear whorls with prominent,
narrow, protractively sinuated radial ribs, 82 on the body whorl,
whose interstices are 2-4 times their width. Microsculpture of very
fine radial riblets, 4-8 between each pair of major ribs, crossed by
barely visible spiral riblets with a prominent secondary sculpture of
closely spaced spiral cording. Sutures moderately impressed, whorls
slightly flattened laterally above and below periphery with evenly
rounded outer margins. Color light yellowish white with irregular
somewhat vague reddish flammulations. Umbilicus V-shaped, mod-
erately open, last whorl decoiling a little more rapidly, contained
5.05 times in the diameter. Aperture subcircular, somewhat flat-
tened basally, inclined about 20 from shell axis. Height of lectotype
1.61 mm., diameter 3.00 mm.
Lectotype. Tonga: Vava'u. Collected by E.
Graeffe. Zoologisches Institut der Universitat, Zurich.
Range. Futuna, Hoorn Islands; Vava'u, Tonga-
tapu, Eua, and Ha'apai Group, Tonga Islands.
Material. Tonga: Vava'u (3 specimens, Zurich):
Utula'aina (Station T-13) near Holonga in heavy forest
at 10-100 ft. elevation (265 specimens, FMNH
152372-3); 200 yd. inland on cliffs at 200-350 ft. eleva-
tion, north side of island (4 specimens, BPBM 87859).
Ha'apai Group, Hufena (= Uoleva) (4 specimens,
Zurich). Eua (6 specimens, Zurich, FMNH 150773,
BMNH 93.9.25.81): heavy primary forest near south
end (Stations T-21, T-22) at 850-1,000 ft. elevation (5
specimens, FMNH 152519, FMNH 152379). Ton-
gatapu: Manima Id., 4 miles northeast of Nuku'alofa
(Station T-4) in second growth on a coral islet (1 speci-
men, FMNH 152397); about 11 miles northwest of
Nuku'alofa, 200 yd. inland (Station T-26) in coastal
scrub (32 specimens, FMNH 152312, FMNH 152461).
Fukave, near Tonga tapu (1 specimen, BPBM 53524).
Hoorn Islands: Futuna (6 specimens, Zurich,
FMNH 150772).
Remarks. Recorded variation between sets of
Sinployea vicaria (table XXVIII) is quite large, with a
spread of 0.75 mm. between the smallest in diameter
(recent collection from Eua) and the largest (type series
from Vava'u). Two factors reduce the possibility that
this has any systematic significance. The specimens in
the Mousson collection at Zurich were selected by
SYSTEMATIC REVIEW
135
TABLE XXVIII. - LOCAL VARIATION IN SINPLOYEA VICARIA.
vicarla vlcarla
NUMBER OF
SPECIMENS
3
RIBS
100.39.21
(82-111)
RIBS/MM.
10.5410.920
(8.72-11.61)
HEIGHT
1.7210.083
(1.62-1.89)
DIAMETER
3.0510.051
(2.98-3.15)
H/D
.56610
(0
RATIO
.034
.526-0.633)
Vavau
Zurich
Hufena, Haapaii
Zurich
4
101.02.68
(95-108)
11.9910.
(11
307
.49-12.82)
1.6010.
(1.
045
49-1.71)
2.7010
(2
.035
.65-2.80)
0.593*0
(0,
,0115
.563-0.612)
Tongatapu 15
Sta. T-26
FMNH 152312, -461
118.33.87
(96-145)
13.340.
(10
369
.59-15.76)
1.5310,
(1,
039
30-1.78)
2.8210
(2
.050
.52-3.09)
.54110
(0,
.0087
,469-0.596)
Fukave, Tongatapu
BPBM 53524
1
128
14.40
1.51
2.83
.535
Eua
Zurich
5
105.012.98
(96-113)
11.8410.
(10
381
.80-13.17)
1.5010.
(1.
034
.42-1.62)
2.8410.049
(2.75-3.03)
.52710.0063
(0.511-0.547)
Sta. T-21
FMNH 152519
4
92.011.53
(90-95)
12.7210.
(11
683
.36-13.42)
1.2510.062
(1.17-1.43)
2.3010
(2
.088
.14-2.55)
.546+0
(0,
,0166
,518-0.585)
BMNH 93.9.25.81
1
143
15.91
1.74
2.86
.609
Put una
Zurich
5
96.2*6.40
(77-111)
12.2010.
(11
442
.03-13.69)
1.4210
(1
.055
.33-1.61)
2.5010
(2
.094
.22-2.80)
.56910
(0
.0100
.546-0.600)
vicaria paucicosta
Vavau
Sta. T-8
FMNH 152488
10
78.015.85
(51-98)
9.3910.
(6.
590
71-11.42)
1.5110
(1
.041
,27-1.78)
2.6310
(2
.038
.42-2.76)
.57410
(0
.012
.500-0.643)
Sta. T-9
FMNH 152414
28
76.811.68
(67-89)
9.9510.210
(8.54-11.44)
1.4310
(1.
,018
,18-1.63)
2.4410
(2
.021
.22-2.62)
.58910
(0
.0062
.518-0.659)
Sta. T-13
FMNH 152372-3
60
89.6*1.20
(66-108)
10.39-10.
(8.
131
08-12.59)
1.48+0
a
,016
,25-1.81)
2.76+0.025
(2.43-3.35)
.536+0.0032
(0.467-0.588)
WHORLS
v. vicaria
Vavau 4 3/8+(4 1/4-4 3/4)
Hufena 4 3/8+(4 1/4-4 5/8)
Tongatapu 4 1/4-OV-4 1/2)
Fukave 4 1/8
Eua
Zurich 4 l/8+(4-4 1/4)
Sta.T-21 4-(3 3/4-4 1/8)
BMNH 41/4
Futuna 4 l/4+(4-4 1/2)
v.paucicos.
Sta. T-8 4 l/2+(4 1/2-5)
Sta. T-9 4 3/8(4 1/4-4 5/8)
Sta. T-13 4 l/4-(3 3/4-4 5/8)
D/U RATIO
SPIRE
APICAL CORDS PROTRUSION
SP/BWW
0.6510
(0
.040
,59-0.72)
4.7110
(4
.312
.09-5.
05)
11
0.2510
CO
.067
.16-0.38)
0.9710
(0
.025
.94-1.02)
0.257*0
(0
.074
.161-0.404)
0.6210,
(0,
,024
,56-0.67)
4.3310
(4
.130
.13-4.
71)
10.33*0.883
(9-12)
0.280
(0
.032
.20-0.35)
0.85*0
(0
.022
.81-0.89)
0.33510
(0
.038
.222-0.389)
0.66+0
(0
,017
,51-0.76)
4.28+0
(3
.092
.88-5.
26)
11.20.374
(10-12)
0.23*0
(0
.014
.16-0.33)
0.86*0.013
(0.77-0.92)
0.26910
(0
.016
.189-0.417)
0.66
4.30
0.20
0.82
0.240
0.63*0
(0
,024
.56-0.61)
4.5210
(4
.116
.20-4.
88)
11.0010.578
(10-12)
0.150
(0
.078
.10-0.21)
0.9110
(0
.021
.87-0.99)
0.167*0
(0
.026
.111-0.241)
0.4710.037
(0.41-0.57)
0.49
0.38+0.038
(0.44-0.66)
4.9110
(4
5.80
4.3410
(4
.224
.43-5.
.172
.03-5.
48)
00)
9.7510.479
(9-11)
0.1710
(0
.012
,15-0.20)
0.72*0
(0
.022
.69-0.79)
0.22810
(0
.0172
.208-0.279)
0.19*0
(0
.022
.13-0.26)
0.84*0
(0
.009
.82-0.87)
0.23110
(0
.0199
.157-0.302)
0.6510.014
(0.59-0.72)
4.06*0
(3
.089
.62-4.
54)
11.2510.479
(10-12)
0.23*0
(0,
.020
.15-0.33)
0.8310
(0
,017
,76-0.89)
0.27610
(0,
,0243
,196-0.385)
0.59+0
(0
.008
.49-0.67)
4.17+0
(3
.042
.73-4.
64)
11.67*0.211
(11-12)
0.21*0.010
(0.10-0.33)
0.7710.
(0,
,007
,69-0.86)
0.265*0,
(0,
,130
,125-0.360)
0.5910.008
(0.46-0.72)
4.7110.054
(3.90-5.
71)
0.16*0,
(0,
.006
07-0.26)
0.9110,
(0,
008
79-1.08)
0.18110,
(0,
0071
080-0.286)
Mousson from field collections that were then depos-
ited in the Godeffroy collection. Presumably these
large specimens can be equated with the retention by
Garrett of larger examples in his collection (for exam-
ple see S. proximo, p. 108). Hence, a major part of the
larger size shown by the Mousson Eua Island and
Vava'u material could result from biased sampling.
Second, the material collected by Price in 1966 was
taken after an extended period of drought. Quite possi-
bly, ecophenotypic dwarfing was involved in the rela-
tively small size of these shells. Since the proportions,
whorl counts, and ribbing are essentially identical, I
am ignoring the size differences and consider them
conspecific. Inclusion of the Futuna shells is less cer-
tain, but their proportions and ribbing match the Ton-
gan examples.
Comparison of measurements in Tables XIX and
XXVII demonstrates that Sinployea vicaria and S.
atiensis are extremely similar in size and proportions.
Direct comparisons of specimens show that in S. vi-
136
SOLEM: ENDODONTOID LAND SNAILS
FIG. 58. a-c, Sinployea vicaria vicaria (Mousson). Vava'u, Tonga. Lectotype. Zoologisches Institut der Universitat Zurich; d, Sinployea
vicaria paucicosta, new subspecies. Station T-9, Mo'ungalafa, Vava'u, Tonga. Holotype. FMNH 158449. Scale lines equal 1 mm. (SH).
caria the secondary spiral cording is much more
crowded, the apical cords reduced in prominence, the
body whorl less distinctly flattened laterally above the
periphery, and the last whorl of the umbilicus decoils
slightly more rapidly. On the average, specimens of S.
vicaria have about one more rib/mm, and perhaps 15
more ribs on the body whorl, but overlap is complete
and these characters cannot be used to separate the
taxa. The similarities are considered to be convergent
and caused by the limited patterns of coiling available
to these snails.
Mousson's original measurements of 3 mm. in
diameter and 4 l /2 whorls are most closely matched by
the specimen from Vava'u selected as lectotype. It is
atypical in having an umbilicus that is slightly nar-
rower than usual because of regular last whorl decoil-
ing (fig. 58c).
Description of soft parts. Foot and tail slightly shorter than
shell diameter, truncated anteriorly, bluntly rounded, and slightly
tapering posteriorly. Sole undivided, pedal grooves prominent, high
on foot, no caudal horn or middorsal groove developed. Slime net-
work typical. Head projecting in front of foot, ommatophores long,
dark in color. Gonopore below right ommatophore and behind right
rhinophore, a vertical, short slit in body wall.
Body color yellow-white on sides and sole of foot, back of head
and neck, ommatophores, and inner part of mantle collar heavily to
moderately speckled with gray.
Mantle collar (MC) wide, white on outer edge, heavily marked
with gray on inner edge, no glandular extension onto pallial roof.
Anus (A) opening just anterior to external ureteric pore (KX).
Pallial region (fig. 60a) extending slightly less than Vfe whorl
apically. Lung roof clear, without granulations. Kidney (K) about
1.45 mm. long, rectal lobe slightly longer than pericardial. Rectal
arm crossing hindgut and extending well down onto parietal wall.
Ureter (KD) typical, a strip of lung roof visible between arms. Heart
(H) nearly parallel to hindgut, % length of pericardial kidney lobe.
Principal pulmonary vein (HV) narrow, unbranched, fading out just
before mantle collar. Hindgut (HG) extending about 0.3 mm. above
kidney apex before starting to deflect downward.
Ovotestis (fig. 60b) a single clump of palmately clavate alveoli
above stomach apex, lying parallel to plane of coiling, sharply trun-
cate anteriorly, soft parts extending one full whorl above ovotestis
apex. Hermaphroditic duct (GD) typical. Albumen gland, talon, pros-
tate, and uterus without unusual features.
Vas deferens (fig. 60c, VD) typical, epiphallus (E) more than half
length of penis, entering penis apically through a longitudinal
epiphallic pore (EP). Penial retractor (PR) short, attached in U-fan to
head of penis, arising from diaphragm. Penis (P) about 1.4 mm. long,
lower third very narrow, internally (fig. 60d) with distinct collar
(PVS) around prominent vergic papilla (PV). Normal circular muscle
collar reduced to a low ridge, main stimulatory pad quite prominent.
Atrium (Y) long, medially swollen.
Free oviduct (UV) enlarging rapidly in diameter just before
junction with spermatheca. Spermatheca (S) and vagina (V) typical.
(Based on FMNH 152373, several expanded adults, 2.73-2.93
mm. in diameter, with 4'/8-4V4 whorls.)
SYSTEMATIC REVIEW
137
FIG. 59. Shell sculpture details of Sinployea uicaria vicaria (Mous-
son). Station T-13, Vava'u, Tonga. FMNH 152373: a, sculpture of
body whorl (535x); b, fracture section through body whorl showing
periostracal (upper) and calcareous (lower) elements of sculpture
(950x).
Sinployea vicaria paucicosta, new subspecies. Fig-
ure 58d.
Diagnosis. Shell small, diameter 2.22-2.76 mm. (mean 2.49
mm.), with 4V4-5 rather tightly coiled whorls. Apex and first post-
nuclear whorl barely emergent, lower whorls descending more
rapidly, body whorl much more rapidly, spire protrusion slightly
more than Vt body whorl width, H/D ratio 0.500-0.659 (mean 0.585).
Apical sculpture of 10-12 (mean 11.5) fine spiral cords. Postnuclear
whorls with prominent, narrow, rather widely spaced, protractively
sinuated radial ribs, 51-98 (mean 77.1) on the body whorl, whose
interstices are 5-8 times their width. Ribs/mm. 6.71-11.44 (mean
9.80). Microsculpture of very fine radial riblets, 812 between each
pair of major ribs, barely visible microspiral riblets, and prominent,
relatively crowded secondary spiral cords. Umbilicus open, U-
shaped, last whorl decoiling more rapidly, contained 3.62-4.64 times
(mean 4.14) in the diameter, margins rounded. Whorl contour and
sutures as in the nominate race.
Sinployea vicaria paucicosta (fig. 58d) differs from
the nominate race in its more sharply defined and more
widely spaced ribbing. The smaller size, slightly
greater H/D ratio, wider umbilicus, and average of one
more apical cord are of uncertain significance.
Sinployea ellicensis (fig. 62a-0 has much more
crowded ribbing and fewer whorls, whereas S.
pseudouicaria (fig. 61a-c) is widely umbilicated and
much larger.
Description. Shell small, with 4% rather tightly coiled whorls.
Apex barely elevated, later whorls descending much more rapidly,
H/D ratio 0.595. Apical whorls and early spire with sculpture eroded.
Postnuclear whorls with narrow, prominent, strongly protractively
sinuated radial ribs, 69 on the body whorl, whose interstices are 4-6
times their width. Microsculpture of very fine radial riblets, 6-12
between each pair of major ribs, crossed by barely visible spiral rib-
lets. Secondary spiral cording prominent and relatively crowded. Su-
tures impressed, whorls strongly rounded above, slightly flattened
laterally above periphery and on basal margin. Color light yellow-
horn, with broad, irregular, reddish flammulations that fade out on
shell base. Umbilicus open, U-shaped, last whorl decoiling more
rapidly, contained 4.00 times in the diameter. Aperture subcircular,
slightly flattened laterally above periphery, inclined about 20 from
shell axis. Height of holotype 1.45 mm., diameter 2.43 mm.
Holotype. Tonga: Vava'u, Station T-9, Mo'un-
galafa, near Tu'anuku, about 14 miles southwest of
Neiafu, at 700 ft. elevation. Collected dead in leaf
mould from patches of heavy forest by Laurie Price on
January 3, 1966. FMNH 158449.
Range. Southern part of Vava'u, Tonga Islands.
Paratypes. Vava'u: near Toulo (Station T-8), 2
miles south of Neiafu in leaf mould in coral rubble
under trees above strandline (16 specimens, FMNH
152488); Mo'ungalafa, near Tu'anuku (Station T-9),
about 14 miles southwest of Neiafu in leaf mould from
heavy forest at 700 ft. elevation (69 specimens, FMNH
152414); Muitola, 15 miles west of Neiafu (Station
T-10) at 400 ft. elevation in heavy forest (3 specimens,
FMNH 158455); cliffs at end of peninsula past Makave
Village (Station T-12), 2 x /2 miles southeast of Neiafu at
200 ft. elevation from leaf mould (1 specimen, FMNH
152360).
Remarks. The four localities for Sinployea ui-
caria paucicosta are all on the south and southeast
coasts of Vava'u. Material of the nominate race is
known on Vava'u from the original collection of
Graeffe in the 1860s, 265 shells from near Holonga
(Station T-13) on the north tip, and four juvenile
specimens from the north side (BPBM 87860) collected
at 200-350 ft. elevation by J. E. Hoffmeister on July
10, 1928. It is probable that they are geographic races,
but the possibility does exist that the variation is
ecophenotypic or that they are separate species. Unfor-
tunately, weather conditions were so dry that no living
material could be obtained of S. v. paucicosta (see also
p. 183).
Material from Stations T-8 and T-9 differs slightly
in size (table XXVIII) but is almost identical in propor-
tions and ribbing. Three examples from Station T-10
are subadult. The single shell from Station T-12 is
badly worn and quite large, diameter 3.13 mm. One
portion of the shell contains traces of widely spaced
ribbing in the suture. Although it has been referred to
this subspecies, the measurements have not been used
in figuring means and ranges because it might belong
to the nominate race. Further collecting is needed in
order to establish the identity of this population.
138
SOLEM: ENDODONTOID LAND SNAILS
The name paucicosta refers to the reduced number
of major radial ribs that characterize this taxon.
Sinployea rotumana (E. A. Smith, 1897). Figure
61d-f.
Champa rotumana E. A. Smith, 1897, Ann. Mag. Nat. Hist., (6) 20,
p. 520 Rotuma Island; Hedley, 1899, Mem. Australian Mus., 3
(7), p. 488 as a synonym of "Charopa modicella" I =Sinployea
ellicensis) from Funafuti.
Diagnosis. Shell of average size, diameter 2.60-3.24 mm.
(mean 2.84 mm.) with 4-4% normally coiled whorls. Apex and spire
slightly to moderately and evenly elevated, last whorl descending
more rapidly, spire protrusion about Vs-'A body whorl width, H/D
ratio 0.481-0.607 (mean 0.547). Apical sculpture of 8-11 (mean 9.67)
fine spiral cords. Postnuclear sculpture of low, rounded, rather
crowded, strongly protractively sinuated radial ribs, 84-144 (mean
105.2) on the body whorl, whose interstices are l'/i-3 times their
width. Ribs/mm. 9.79-14.15 (mean 1L74). Microsculpture of fine ra-
dial riblets, 4-7 between each pair of major ribs, slightly finer and
more crowded spiral riblets, with secondary spiral cording that be-
comes extremely fine on lower spire and very crowded. Umbilicus
open, U-shaped, last whorl decoiling slightly more rapidly, contained
3.93-4.86 times (mean 4.32) in the diameter, margins rounded. Su-
tures deep, whorls rounded to strongly compressed laterally above
periphery, aperture inclined about 20 from shell axis.
Sinployea rotumana (fig. 61d-f) has the major rib-
bing lower and less prominent, the secondary spiral
cording much more crowded, and the body whorl more
strongly flattened laterally than in S. vicaria (fig.
58a-c). Sinployea atiensis (fig. 44d-f) from the Cook
Islands has fewer major radial ribs, more apical cords,
and much more widely spaced secondary spiral cord-
ing. There are also subtle differences in umbilical and
whorl contours.
Description. Shell of average size, with a little more than 4
normally coiled whorls. Apex and early spire barely elevated, body
LM
PR
PVS
EP
FIG. 60. Anatomy of the Tongan Sinployea vicaria vicaria (Mousson). Station T-13, near Holonga, Vava'u, Tonga Islands. FMNH 152373: a,
pallial region; b, apical genitalia; c, terminal genitalia; d, interior of penis. Scale lines equal 1 mm. (CW).
a
^^^_ ..'-J"/,ti^
FIG. 61. a-c, Sinployea pseudovicaria, new species. Vaitupu, Ellice Islands. Holotype. BPBM 189682; d-f, Sinployea rotumana (E. A.
Smith). Rotuma Island. BPBM 187049. Scale lines equal 1 mm. (SH).
139
140
SOLEM: ENDODONTOID LAND SNAILS
whorl descending a little more rapidly, H/D ratio 0.497. Apical
whorls 1%, sculpture of 10 prominent spiral cords, with a very faint
secondary sculpture of radial riblets on last quarter whorl. Postnu-
clear whorls with prominent, rounded, low, slightly protractively
sinuated radial ribs, 87 on the body whorl, whose interstices are 2-3
times their width. Microsculpture of fine radial riblets crossed by
exceedingly fine spiral riblets with a secondary sculpture of minor,
very crowded spiral cords. Sutures deep, whorls strongly rounded
above, almost evenly rounded on outer and basal margins. Color
light yellowish horn with broad, irregular reddish flammulations
coalescing on base of shell. Umbilicus narrow, U-shaped, regularly
decoiling, contained 4.22 times in the diameter. Aperture subcircu-
lar, slightly flattened laterally above periphery, inclined about 20
from the shell axis. Height of lectotype 1.38 mm., diameter 2.78 mm.
Lectotype. Rotuma Island. Collected by J. Stan-
ley Gardiner. BMNH 77.11.8.57.
Range. Rotuma Island, 300 miles northwest of
Fiji.
Parades. BMNH 77.11.8.58-63.
Material. Rotuma (1 specimen, BPBM 187049).
Remarks. The single specimen collected by
Harold St. John in 1938 (BPBM 187049) has the body
whorl more compressed laterally above the periphery,
but otherwise compares well with the type set.
Differences from Sinployea vicaria and S. atiensis are
covered in the diagnosis above.
Sinployea ellicensis, new species.
Materials from Funafuti and Nukulaelae collected
by Graeffe in 1868 and Charles Hedley in 1899 differ
from the Tongan S. vicaria (fig. 58a-d) in their
stronger ribbing and nearly circular aperture. Species
of similar size from Eastern Polynesia have the aper-
ture strongly flattened laterally above the periphery
and usually show large differences in sculpture and/or
shape. Species from Fiji, such as S. godeffroyana (fig.
69a-c) and S. princei (fig. 65a-c), differ in ribbing and
higher spire (latter) or umbilical shape and much finer
ribbing (former).
Recognition of two subspecies is done with some
hesitation because only a single collection has been
made on Nukulaelae. Specimens from this island are
much smaller and have more apical cords than the
examples from Funafuti. Although an equally great
size difference is found between populations of
Sinployea vicaria on Tongatapu and Eua Islands (table
XXVIII), the distinct difference in island topography
between the latter islands suggests that the distinc-
tions may be environmentally correlated. Both
Funafuti and Nukulaelae are atolls. Presumably there
is no major habitat difference involved.
Sinployea ellicensis ellicensis, new species and sub-
species. Figure 62a-c.
Patula vicaria Mousson, 1873 (not Mousson, 1871), J. de Conchyl.,
21, p. 104 Funafuti, Ellice Islands.
Champa modicella Hedley, 1899 (not Ferussac, 1840), Mem. Aus-
tralian Mus., 3 (7), p. 488 Funafuti, Ellice Islands.
Diagnosis. Shell small, diameter 2.20-2.81 mm. (mean 2.47
mm.), with 4-4M> rather tightly coiled whorls. Apex and spire
slightly to moderately and evenly elevated, body whorl descending
more rapidly, spire protrusion about Vs-Vs body whorl width, H/D
ratio 0.496-0.561 (mean 0.549). Apical sculpture of 8-12 (mean 9.42)
prominent spiral cords. Postnuclear whorls with prominent,
rounded, crowded, slightly protractively sinuated radial ribs, 79-125
(mean 100.3) on the body whorl, whose interstices are usually less
than twice their width. Ribs/mm. 10.55-16.69 (mean 13.23). Micro-
sculpture of fine radial riblets, 3-5 between each pair of major ribs,
barely visible and extremely crowded spiral riblets, with a secondary
microsculpture of spiral cords slightly larger and more widely spaced
than the microradials. Umbilicus open, U-shaped, last whorl decoil-
ing more rapidly, contained 3.48-5.46 times (mean 4.39) in the
diameter, margins rounded. Whorls strongly and evenly rounded,
aperture circular, not or only slightly flattened laterally above
periphery, inclined about 20 from shell axis.
Sinployea ellicensis (fig. 62a-c) differs from the
Tongan S. vicaria (fig. 58a-c) in its smaller size, lower
spire protrusion, larger radial ribs, and more rounded
whorl contours. Sinployea ellicensis nukulaelaeana
(fig. 62d-f) is much smaller and has more apical cords.
Sinployea princei (fig. 65a-c) has more whorls, mono-
chromatic coloration, and only traces of secondary spi-
ral cording.
Description. Shell smaller than average, with 4% normally
coiled whorls. Apex and spire moderately and evenly elevated, body
whorl descending much more rapidly, H/D ratio 0.556. Apical whorls
1%, sculpture of 10 prominent, relatively widely spaced spiral cords.
Postnuclear whorls with rounded, relatively crowded, protractively
sinuated radial ribs, 119 on the body whorl, whose interstices are 2-3
times their width. Microsculpture of fine radial riblets, 3-6 between
each pair of major ribs, crossed by barely visible spiral riblets and
rather crowded secondary spiral cords. Sutures deeply impressed,
whorls strongly rounded above, very slightly flattened laterally
above periphery. Umbilicus narrowly U-shaped, last whorl decoiling
more rapidly, contained 3.89 times in the diameter. Color light yel-
lowish horn with vague, irregular reddish flammulations, fainter on
base of shell. Aperture circular, slightly flattened laterally above
periphery, inclined about 10 from the shell axis. Height of holotype
1.56 mm., diameter 2.81 mm.
Holotype. Ellice Islands: Funafuti. Collected by
Charles Hedley. AMS C.6124.
Range. Funafuti, Ellice Islands.
Paratypes. Ellice Islands (3 specimens, ex BPBM
1667): Funafuti (102 specimens, AMS C.6124, Zurich,
DMW MF-8810, FMNH 116981, FMNH 158451).
Remarks. Adult specimens in the Mousson col-
lection are distinctly higher and slightly larger (table
XXIX) than those collected by Hedley (AMS C.6124).
Probably this is another example of collection bias to-
ward retention of larger individuals. Differences from
Sinployea vicaria are covered in the diagnosis above.
Although direct comparison of examples leaves no
doubt that they are distinct, measurements do overlap
extensively. Unlocalized specimens would be hard to
identify without direct comparisons. Measurements of
S. princei (Liardet) are almost identical (table XXX),
but there is a distinct difference in coiling pattern.
Sinployea ellicensis nukulaelaeana, new sub-
species. Figure 62d-f.
Diagnosis. Shell very small, diameter 1.99-2.20 mm. (mean
2.08 mm.), with 3%-4 normally coiled whorls. Apex and spire
slightly and evenly elevated, last whorl descending more rapidly,
spire protrusion about Ve body whorl width, H/D ratio 0.496-0.561
(mean 0.533). Apical sculpture of 11-13 (mean 11.67) prominent spi
a
FIG. 62. a-c, Sinployea ellicensis ellicensis, new species and subspecies. Funafuti, Ellice Islands. Holotype. AMS C6124; d-f, Sinployea
ellicensis nukulaelaeana, new subspecies. Nukulaelae, Ellice Islands. Holotype. AMS C6123. Scale lines equal 1 mm. (SH).
141
142
SOLEM: ENDODONTOID LAND SNAILS
TABLE XXIX. - LOCAL VARIATION IN WESTERN POLYNESIAN SINPLOYEA.
NUMBER OF
SPECIMENS RIBS
RIBS/MM.
ellicensls
ellicensis
Zurich
10 104.1*3.16
(91-125)
13.380.474
(11.02-16.71)
AMS C6124
21 98.12.33
(79-119)
13.230.238
(10.55-16.69)
ellicensis
nukulaelaeana
AMS C6123
6 84.5*4.85
(72-102)
13.02+0.734
(11.08-15.20)
Vaitupu
BPBM 1667
5 113.54.51
(109-118)
11.15*0.156
(10.99-11.30)
Zurich
3 111.3*4.34
(107-120)
10.81*0.020
(10.79-10.85)
Eua
BPBM 7983
2 87.0*6.02
(81-93)
8.540.110
(8.43-8.65)
Zurich
3 100.017.51
(87-113)
9.19*0.371
(8.50-9.77)
rotumana
BMNH 77.11.8.57-63
8 104.0*7.01
(84-144)
11.74+0.519
(9.79-14.15)
H/D RATIO
1.42*0.023 2.50*0.028 0.570*0.0066
(1.29-1.51) (2.40-2.65) (0.534-0.596)
1.29*0.022 2.42+0.033 0.5330.0061
(1.14-1.57) (2.22-2.83) (0.486-0.592)
1.11+0.016 2.08+0.032 0. 53310-.0102
(1.06-1.16) (1.99-2.20) (0.496-0.561)
1.55*0.066 3.18+0.050 0.487+0.0152
(1.42-1.76) (3.05-3.34) (0.454-0.525)
1.58+0.029 3.30+0.121 0.479+0.0121
(1.52-1.62) (3.18-3.54) (0.458-0.500)
1.53+0.082
3.24+0.181
(1.45-1.61)
(3.06-3.42)
1.67+0.048 3.45+0.125
(1.58-1.74) (3.26-3.68)
1.560.063
2.86+0.066
(1.26-1.82)
(2.62-3.26)
0.472+0.0001
(0.471-0.473)
0.4830.0157
(0.455-0.509)
0.544+0.0138
(0.481-0.607)
WHORLS
e. ellicensis
Zurich 4 3/8-(4 1/4-4 1/2)
AMS C6124 4 l/8+(4-4 1/2)
e.nukulaelae.
AMS C6123 3 7/8-(3 5/8-4)
pseudovicarla
Vaitupu 4
BPBM 1667
Eua 4
BPBM 7983
Zurich 4
3/8(4 1/4-4 1/2)
l/2+(4 1/2-4 5/8)
3/8(4 1/4-4 1/2)
l/2+(4 1/2-4 3/4)
rotumana
BMNH 4 l/4-(4-4 5/8)
UMBILICUS D/U RATIO APICAL CORDS
0.590.022 4.24+0.117 10.50.77
(0.49-0.76) (3.48-4.86) (9-12)
0.5510.016 4.43+0.081 9.42+0.21
(0.43-0.72) (3.77-5.46) (8-11)
0.48+0.013 4.31+0.086 11.7+0.34
(0.44-0.53) (4.13-4.66) (11-13)
0.95+0.044 3.35+0.197 11.7+0.33
(0.79-1.05) (2.94-4.05) (11-12)
1.12+0.050 2.93+0.034 11
(1.05-1.22) (2.89-3.00)
1.0510.066 3.08+0.020
(0.99-1.12) (3.06-3.10)
1.100.079 3.1710.109 9.510.50
(0.99-1.25) (2.95-3.30) (9-10)
0.65+0.025 4.3510.117 9.67+0.88
(0.58-0.81) (3.93-4.86) (8-11)
ral cords. Postnuclear sculpture of narrow, prominent, crowded, pro-
tractively sinuated radial ribs, 75-102 (mean 84.5) on the body
whorl, whose interstices are l'/2-3 times their width, often with
periostracal extensions. Ribs/mm. 11.06-15.20 (mean 13.02). Micro-
sculpture of fine radial riblets, 3-6 between each pair of major ribs,
crossed by barely visible spiral riblets, with secondary spiral cording
that is rather crowded and equal in spacing to microradials. Um-
bilicus open, U-shaped, last whorl decoiling more rapidly, contained
4.13-4.66 times (mean 4.31) in the diameter, margins rounded.
Aperture and whorls circular, slightly compressed laterally above
periphery.
Sinployea ellicensis nukulaelaeana differs from the
nominate race in possessing more apical spiral cords
and in being much smaller.
Description. Shell very small, with slightly less than 4 nor-
mally coiled whorls. Apex and spire moderately and evenly elevated,
body whorl descending rapidly, H/D ratio 0.547. Apical whorls 1%,
sculpture of 12 prominent spiral cords. Postnuclear whorls with
prominent, narrow, protractively sinuated, crowded, radial ribs, 72
on the body whorl, whose interstices are l'/2-3 times their width.
Microsculpture of fine radial riblets, 3-6 between each pair of major
SYSTEMATIC REVIEW
143
ribs, barely visible spiral riblets, with relatively prominent and
crowded secondary spiral cording. Sutures deep, whorls strongly
rounded above, very slightly compressed laterally above periphery
and on basal margin, with evenly rounded outer margin. Color light
yellow-horn, with vague and irregular reddish flammulations that
tend to coalesce on shell base. Umbilicus open, U-shaped, last whorl
decoiling slightly more rapidly, contained 4.66 times in the diameter,
margins rounded. Aperture circular, slightly flattened laterally
above periphery and on basal margin, inclined about 15 from shell
axis. Height of holotype 1.14 mm., diameter 2.07 mm.
Holotype. Ellice Islands: Nukulaelae. AMS
C.6123.
Range. Nukulaelae, Ellice Islands.
Paratypes. Nukulaelae (7 specimens, AMS
C.6123, FMNH 158452).
Remarks. Size differences between the races of
Sinployea ellicensis (table XXIX) equal those found be-
tween the Tongatapu and Eua Island S. vicaria col-
lected in 1965 (table XXVIII). Subspecific recognition
in this case is based on the difference in apical cord
count and absence of major topographic differences be-
tween Funafuti and Nukulaelae. There are no consis-
tent sculptural or shape differences between the popu-
lations.
contained 2.76 times in the diameter. Aperture circular, slightly flat-
tened above periphery, inclined about 25 from shell axis. Height of
holotype 1.25 mm., diameter 2.73 mm.
Holotype. Ellice Islands: Vaitupu. Collected by L.
Isaacs in May, 1941. BPBM 189682.
Range. Vaitupu, Ellice Islands.
Paratypes. Ellice Islands (5 specimens, BPBM
1667): Vaitupu (8 specimens, BPBM 106246, BPBM
189682, Zurich, FMNH 158450).
Material. "Eua" (error) (7 specimens, BPBM
7983 ex Mus. Godeffroy, FMNH 116980, Zurich ex
Mus. Godeffroy).
Remarks. Although the holotype is subadult, it
comes from the only set whose locality data can be
accepted without question. The specimens from the
Museum Godeffroy labeled "Eua" are essentially iden-
tical to the Vaitupu shells (table XXIX) and probably
were accidentally mislabeled. Nothing at all similar
has been recorded from Eua subsequently.
In view of the confusion between this species and
vicaria in the older literature, I have chosen to use the
name pseudovicaria.
Sinployea pseudovicaria, new species. Figure 61a-c.
Patula vicaria var. major Schmeltz, 1869, Cat. Mus. Godeffroy, 4,
p. 72 Vaitupu, Ellice Islands (nude name); Schmeltz, 1874,
Cat. Mus. Godeffroy, 5, p. 93 (nude name).
Patula modicella Mousson, 1873 (not Ferussac, 1840), J. de
Conchyl., 21, pp. 104-105 Vaitupu, Ellice Islands.
Diagnosis. Shell relatively large, diameter 3.03-3.52 mm.
(mean 3.24 mm.), with 4'/4-4% normally coiled whorls. Apex and
early spire flat, lower spire descending slightly, body whorl descend-
ing moderately, spire protrusion about Vs body whorl width, H/D
ratio 0.454-0.525 (mean 0.486). Apical sculpture of 10-13 (mean
11.75) fine spiral cords. Postnuclear whorls with fine, prominent,
protractively sinuated radial ribs, 82-127 (mean 108.1) on the body
whorl, whose interstices are 2-4 times their width. Ribs/mm. 9.56-
15.15 (mean 11.33). Microsculpture of prominent radial riblets, 3-6
between each pair of major ribs, crossed by extremely fine and
crowded spiral riblets. No secondary spiral cording present. Um-
bilicus broadly open, regularly decoiling, cup-shaped, contained
2.89-4.05 times (mean 3.17) in the diameter, margins rounded.
Whorls with subcircular outline, very slightly flattened laterally
above periphery, inclined about 25 from shell axis.
The flat spire, widely open umbilicus, and larger
size immediately separate Sinployea pseudovicaria
from the other Ellice and Tongan species. Sinployea
lamellicosta (fig. 40a-c) from Tahiti has an equally
wide umbilicus and is nearly identical in size and pro-
portions, but has comparatively few radial ribs, a de-
pressed or flat spire, and fewer, more loosely coiled
whorls.
Description. Shell rather small, with 4V4 relatively loosely
coiled whorls. Apex and early spire flat, last whorl barely descend-
ing, H/D ratio 0.458. Apical whorls 1%, sculpture of 13 moderately
prominent spiral cords. Postnuclear whorls with prominent,
rounded, protractively sinuated radial ribs, 82 on the body whorl,
whose interstices are 2-4 times their width. Microsculpture of fine,
crowded radial riblets with much finer and more crowded spiral rib-
lets. Sutures deeply impressed, whorls slightly shouldered above,
body whorl slightly flattened laterally above periphery. Color par-
tially leached from shell with dark, reddish brown flammulations
remaining. Umbilicus broadly open, U-shaped, regularly decoiling,
Sinployea kusaieana, new species. Figures 63a-c,
67h.
Diagnosis. Shell smaller than average, diameter 2.52-2.75
mm. (mean 2.60 mm.) with 3%-4'/8 normally coiled whorls. Apex and
spire almost flat, slightly and evenly to moderately elevated, last
whorl descending somewhat to much more rapidly, spire protrusion
about V? body whorl width, H/D ratio 0.462-0.542 (mean 0.506). Api-
cal sculpture of 8-10 (mean 9.20) fine, rather inconspicuous spiral
cords, whose interstices are 3-4 times their width. Postnuclear
sculpture of prominent, strongly protractively sinuated, rather
crowded, V-shaped radial ribs, 84-103 (mean 91.1) on the body
whorl, whose interstices are 3-5 times their width. Microsculpture of
very fine radial riblets, 5-10 between each pair of major ribs, crossed
by slightly finer and more crowded spiral riblets, with secondary
spiral cording slightly more prominent than microradials, whose in-
terstices are about 3 times their width. Umbilicus broadly open, V-
shaped, last whorl usually decoiling more rapidly, contained 3.21-
3.67 times (mean 3.51) in the diameter, margins rounded. Suture
deep, whorls strongly rounded above, flattened laterally above
periphery and weakly flattened in columellar margin. Aperture sub-
circular, flattened laterally above periphery, inclined about 20 from
shell axis. Color reddish yellow-horn, without darker markings.
Sinployea kusaieana (fig. 63a-c) differs from the
other barrier-free Caroline Island species by its posses-
sion of an epiphallus, vergic papilla, and Sinployea-
type stimulatory pad in the penis, plus having the kid-
ney arms equal in length. Trukcharopa trukana (fig.
63d- f) is dark reddish brown in color and has an aver-
age of 150 major ribs on the body whorl but is virtually
identical in many measurements (pp. 209-210). Rus-
satus nigrescens (fig. 89d-f) is much more loosely coiled
and very much larger in size with fewer whorls and a
very narrow umbilicus. Kubaryiellus kubaryi (fig.
89a-c) has a depressed apex and is much larger. The
most similar Polynesian species areS. navutuensis (fig.
68d-f) from the Lau Archipelago, which has 127-147
ribs on the body whorl and averages one-quarter whorl
less, and S. rotumana (fig. 61d-f) and S. vicaria (fig.
a-f
Fig. 63. a-c, Sinployea kusaieana, new species. Station 44, Lele Islet, Kusaie, Caroline Islands. Holotype. BPBM 155874; d-f, Trukcharopa
trukana, new species. Station A5, Mt. Urupot, Tol Islet, Truk, Caroline Islands. Holotype. BPBM 189318. Scale line equals 1 mm. (a-c, e, SH; d, f,
MM).
144
SYSTEMATIC REVIEW
145
58a-c), which have higher spires and narrower um-
bilici.
Description. Shell smaller than average, with 4 moderately
tightly coiled whorls. Apex flat, spire and body whorl descending
slightly, H/D ratio 0.481. Embryonic whorls l'/2, sculpture of 9 nar-
row, relatively low spiral ribs. Remaining whorls with sculpture of
crowded, rounded, protractively sinuated radial ribs, 84 on the body
whorl, whose interstices are 2-3 times their width. Microsculpture of
fine radial riblets, 5-10 between each pair of major ribs, crossed by
finer, slightly more crowded spiral ribs, with more prominent,
rounded, more widely spaced spiral cords. Sutures deeply impressed,
whorls sharply rounded above, somewhat flattened laterally with
slightly flattened columellar margin. Umbilicus broadly V-shaped,
regularly decoiling, contained 3.44 times in the diameter. Color light
reddish yellow-horn without traces of flammulations. Aperture sub-
circular, strongly flattened laterally above periphery with slightly
flattened columellar margin, inclined about 15 from the shell axis.
Height of holotype 1.25 mm., diameter 2.60 mm.
Holotype. Caroline Islands: Kusaie, Station 44,
Lele Islet, under stones and logs at 3 ft. elevation. Col-
lected by S. Ito and Yoshio Kondo on January 21, 1936.
BPBM 155874.
Range. Kusaie, Caroline Islands.
Paratypes. Kusaie: Lele Islet (Station 44) under
stones and logs at 3 ft. elevation (59 specimens, BPBM
155874-7); Mot District (Station 66) 300 yd. west of
Kefwas, 10 yd. inland at 4-5 ft. elevation (1 specimen,
BPBM 156179).
Remarks. Both of the known localities on Kusaie
are lowland elevations under disturbed conditions.
Quite possibly Sinployea kusaieana will be found to
occur on other of the Eastern Caroline Islands.
Description of soft parts. Foot and tail equal to shell diameter,
tapering posteriorly. Sole and pedal grooves typical. Gonopore in
normal position.
Body color light yellow-white, without darker markings.
Pallial region extending % whorl apically, without unusual fea-
tures. Lung roof clear, no granulations. Kidney typically bilobed,
maximum length 1.0-1.3 mm., lobes equal in length. Ureter typical,
narrow strip of lung roof visible between arms.
Ovotestis, hermaphroditic duct, albumen gland, talon, prostate
(fig. 67h, DG), and uterus typical, no unusual features.
Vas deferens (VD) not expanded in any area, entering epiphallus
through usual pore. Epiphallus (E) about half as long as penis, slen-
der, only weakly expanded. Penial retractor (PR) short, arising from
diaphragm, inserting on head of penis lateral to epiphallic junction.
Penis (P) about 1 mm. long, quite slender, upper half weakly ex-
panded, internally with typical vergic papilla and small stimulatory
pad. Atrium (Y) long and slender.
Free oviduct (UV) typical. Spermatheca (S) with basal portion of
shaft less expanded than usual. Vagina (V) tapering to atrium.
(Based on BPBM 155874, 4 whole and several partial speci-
mens.)
Sinployea sp.
Material. Marianas: Saipan, north edge of Gara-
pan at base of a century plant (2 specimens, BPBM
190043).
Remarks. The two dead, somewhat worn shells
collected by Harold W. Harry in October 1944 are not
referable to any described species. They are most simi-
lar to Sinployea atiensis (Pease) from the Cook Islands
and S. ellicensis from Funafuti, but are smaller and
with fewer whorls than the former and higher and with
fewer ribs than the latter. The coiling pattern is looser
than in either species.
Because they were collected under an introduced
plant in an urban area, the possibility exists that they
were introduced on plants from Polynesia. Without
additional information, I am not willing to accept this
as a valid range extension and include the specimens
here only to call attention to the need for making more
collections in the Northern Marianas. Measurements
of the shells have been included in Table XXVII for
convenient reference.
FIJIAN Sinployea
There is an extraordinary diversity of Sinployea
and derivative genera in Fiji and the Lau Archipelago
(table XXX). Two species, S. irregularis (Garrett) and
S. godeffroyana, are known from Viti Levu; the former
also lives on Ovalau; and one species, S. princei (Liar-
det), is known from Taveuni and possibly Kandavu.
The absence of any records from relatively large is-
lands such as Vanua Levu, Koro, Ngau, Moala, and
Thithia probably are artifacts of collecting activity. I
know of no significant collections of land mollusks from
any of the above islands. Undoubtedly, most of them
will be found to have Sinployea present. Six species of
Sinployea are known from the Lau Group, one of them,
S. inermis (Mousson), with three geographic races.
Genera that are probably derived directly from
Sinployea include Maafu thaumasius, which is re-
stricted to Nayau in the Lau Group, and Ba humbugi
on Viti Levu. The former agrees in shell micro-
sculpture with Sinployea inermis, although radically
altered in shell form and size (fig. 76a-c), whereas the
latter (fig. 74a-c) can be derived without difficulty
from S. irregularis. I am less certain of the probable
derivation for the Vanua Mbalavu Lauopa
mbalauuana (fig. 77a-c).
In Polynesia it is very unusual to have any sym-
patric occurrences of Sinployea or even for two species
to live in the same valley, but there is extensive sym-
patry in the southern Lau Archipelago. The northern
Lau islands of Kimbombo, Vanua Mbalavu, Munia,
and Mango have only single species records, either S.
inermis or S. adposita. On Nayau, S. adposita and S.
lauensis were taken at the same station, whereas on
Aiwa, Tuvutha, and Lakemba both S. adposita and S.
inermis were found together. On Oneata, both S.
navutuensis and S. adposita were collected, but not at
the same stations. Wangava had large numbers of S.
recursa and a few S. lauensis taken together, whereas
on Navutu-i-Loma H. S. Ladd collected S. inermis, S.
lauensis, and S. navutuensis together at two different
stations. Finally, on Yangasa Levu large numbers of S.
inermis, S. adposita, and S. lauensis were sympatric.
Unfortunately most of these specimens were dead indi-
viduals collected during the July to August dry season.
Soft parts were available only for S. inermis and S.
adposita. Dissection of specimens from Station 97,
146
SOLEM: ENDODONTOID LAND SNAILS
TABLE XXX. - RANGE OF VARIATION IN FIJIAN SINPLOYEA.
NUMBER OF
NAME
SPECIMENS
angularis
188
re curs a
108
prince!
(Liardet)
Taveuni
9
Kandavu
U
inermis inermis
197
(Mousson)
inermis
meridionalis
169
inermis
lakemhana
14
lauensis
82
navutuensis
29
jtdjjp ^s i t a
176
irregularis
78
(Garrett)
godef f royana
RIBS RIBS/MM. HEIGHT DIAMETER H/D RATIO
38.0(36-42) 5.67(5.28-6.16) 1.21(1.08-1.29) 2.11(2.02-2.27) 0.574(0.512-0.615)
REDUCED 1.66(1.51-1.84) 2.83(2.60-3.09) 0.587(0.562-0.634)
91.1(80-102) 12.06(11.84-13.80) 1.34(1.26-1.57) 2.43(2.35-2.68) 0.551(0.524-0.586)
96.3(84-119) 13.10(11.4-14.67) 1.27(1.17-1.45) 2.33(2.02-2.58) 0.545(0.521-0.577)
80.9(65-104) 9.82(8.39-11.75) 1.46(1.18-1.84) 2.60(2.24-3.06) 0.562(0.500-0.629)
72.2(53-104) 8.37(6.75-11.70) 1.64(1.32-1.94) 2.88(2.50-3.22) 0.566(0.505-0.655)
51.3(38-62) 7.01(4.67-8.20) 1.37(1.10-1.64) 2.36(2.14-2.86) 0.583(0.508-0.634)
103.9(95-119) 11.53(10-93-14.46) 1.47(1.18-1.78) 2.76(2.40-3.13) 0.530(0.493-0.581)
136.7(121-147) 17.36(16.94-17.62) 1.32(1.16-1.49) 2.66(2.58-2.78) 0.498(0.499-0.566)
53.3(36-74) 5.28(3.35-7.23) 1.81(1.58-2.20) 3.29(2.70-3.62) 0.603(0.538-0.687)
IRREGULAR 1.77(1.51-2.30) 2.87(2.50-3.52) 0.614(0.528-0.764)
117.7(85-152) 28.36(20.99-38.03) 1.26(1.19-1.29) 2.12(1.95-2.32) 0.595(0.577-0.619)
WHORLS
UMBILICUS
D/U
RATIO
APICAL
CORDS
SPIRE
ELEVATION
BODY WHORL
WIDTH
SP/BWW
angul.
3 7/8+O 3/4-4)
0.35(0.23-0
,40) 6.14(5
.21-8.71)
14.6(13-16)
0.19(0.16-0.23)
0.62(0.
59-0.66)
0.316(0.
250-0.389)
recur.
3 7/8-O 1/2-4 1/8)
0.43(0.33-0
,53) 6.63(5
.75-7.90)
15.9(14-18)
0.23(0.
20-0.30)
0.86(0.
81-0.91)
0.264(0.
222-0.346)
princ.
3 3/4+(3 1/2-4 1/8)
0.57(0.49-0.
63) 4.28(3
.74-4.83)
9(8-10)
0.12(0.
08-0.23)
0.81(0.
70-0.89)
0.151(0.
095-0.259)
Tav.
Kan.
3 5/8+(3 1/2-4)
0.46(0.39-0,
53) 4.76(4
.14-5.19)
NOT
AVAILABLE
0.13(0.
10-0.18)
0.75(0.
66-0.86)
0.179(0.
130-0.250)
Line.
4-(3 1/2-4 5/8)
0.62(0.49-0.
82) 4.17(3
.47-4.69)
8.96(6-11)
0.16(0.
10-0.25)
0.81(0.
72-0.92)
9.192(0.
115-0.280)
i.mer.
4 l/4+(3 7/8-4 5/8)
0.82(0.69-0,
99) 3.51(3
.07-3.81)
11.6(9-14)
0.20(0.
13-0.26)
0.96(0.
82-1.05)
0.207(0.
140-0.285)
l.lak.
4-t-O 3/4-4 1/2)
0.45(0.33-0.76) 5.54(3
.78-7.10)
13.6(11-16)
0.19(0.
13-0.23)
0.82(0.
72-0.89)
0.226(0.
153-0.259)
lauen.
3 7/8(3 1/2-4 1/4)
0.65(0.53-0.
79) 4.24(3
.88-4.65)
9.6(9-11)
0.10(0.
03-0.16)
0.91(0.
82-1.09)
0.113(0.
038-0.161)
navut.
4 1/8(4-4 3/8)
4 3/8+(4-4 7/8)
0.78(0.76-0,
0.28(0.17-0,
82) 3.38(3
46)11.9(7.
.25-3.52)
85-17.0)
10.3(9-11)
13.0(12-14)
0.09(0.
0.24(0.
07-0.13)
13-0.33)
0.74(0.
1.04(0.
66-0.79)
86-1.18)
0.125(0.
0.227(0.
100-0.167)
133-0.275)
adpos.
irreg.
3 3/4+(3 1/4-4 3/8)
0.64(0.39-0,
92) 4.54(3
.79-6.83)
13.5(16-20)
0.20(0.
10-0.46)
0.96(0.
86-1.09)
0.201(0.
100-0.424)
godef.
3 3/4+(3 3/4-4)
0.42(0.33-0,
56) 5.26(4
.11-6.30)
9.7(8-11)
0.12
0.81
0.143
Toumba, Lakemba, Lau, showed that S. adposita had a
longer penis with very small stimulatory pad and nor-
mal vergic papilla, whereas S. inermis had a shorter
penis with very large stimulatory pad and small vergic
papilla. On Viti Levu, S. godeffroyana and S. ir-
regularis have been collected at the same station, and I
have found S. irregularis and Ba humbugi under the
same piece of bark.
Evidence from the white body color suggests that
S. irregularis, S. inermis, and S. adposita are strictly
terrestrial. Generally, semiarboreal species have gray
color on the ommatophores and neck, sometimes also
on the mantle collar edge. Two species, S. angularis
and S. recursa, have greatly reduced sculpture, and I
suspect that they may be semiarboreal or arboreal
species. Unfortunately, they are known only from dead
examples.
Patterns of distribution are relatively simple.
Sinployea irregularis and S. godeffroyana are widely
distributed on forested areas of Viti Levu. Sinployea
princei was described from Taveuni, and essentially
identical specimens coming from the Rolle collection
are labeled Kandavu. Sinployea inermis and S. ad-
posita are widely distributed in the Lau Archipelago,
with the former having a slightly greater range.
Sinployea recursa is known only from Wangava, and S.
angularis, only from Namuka. Sinployea navutuensis
is recorded from Navutu-i-Loma, and a single shell
from Oneata has been referred here with some hesita-
tion. Sinployea lauensis has a fairly wide range in
southern Lau, having been found on Nayau, Wangava,
Namuka, Navutu-i-Loma, and Yangasa Levu.
Many of the Lau Group have never been collected
for land mollusks. Generally, only a single or at most
SYSTEMATIC REVIEW
147
two stations made in a hurried visit of one to two days
account for the collecting done to date on most islands.
Obviously there are splendid opportunities for further
field work. Particularly important will be efforts to
work out patterns of species isolation between sympat-
ric taxa.
Sinployea inermis (Mousson) from the Lau Ar-
chipelago, S. princei (Liardet) from Taveuni and Kan-
davu, and S. irregularis (Garrett) from Viti Levu are
the generalized Fijian species. Sinployea inermis is one
of the most "average" Sinployea, with the monochrome
coloration, details of sculpture, and proportions (fig.
66a-c) differentiating it from the Tongan S. vicaria
(Mousson) (fig. 58a-c) and the Ellice Island S. ellicen-
sis (fig. 62a-c). Subspecies of S. inermis depart further
from the typical pattern, but are still generalized in
shape and form. Sinployea princei (fig. 65a-c) has a
weakly flammulated color pattern, much more crowded
radial ribbing, and reduced secondary spiral cording,
but is very similar to S. inermis. Greater differences
are shown by S. irregularis (fig. 65d-f). The spire is
flat, sutures deep, apical sculpture much finer and
more numerous, ribbing very irregular, body whorl
often abruptly descending, and the size is slightly
larger.
Several species show minor character changes.
Sinployea adposita (Mousson) (fig. 69d-f) can be de-
rived from S. inermis by increase in size, narrowing of
the umbilicus, and increase in rib spacing. The two
species are sympatric and show character displacement
in penial structure, with S. inermis (fig. 67e-f) having
the stimulatory pad and muscular collar proportion-
ately larger than usual and the vergic papilla shorter,
whereas in S. adposita (fig. 67g) the vergic papilla is
quite large. Sinployea lauensis (fig. 68a-c) has more
crowded ribbing, a less protruded spire, fatter body
whorl, and more rapidly decoiling umbilicus. It has not
been dissected. Sinployea navutuensis (fig. 68d-f) has
much more crowded radial ribbing, a wider umbilicus,
nearly flat apex, and lower H/D ratio. Two other Lau
Archipelago species are more strongly differentiated.
The slightly larger S. recursa (fig. 64a-c) has a
strongly flammulated color pattern and the sculpture
greatly reduced. No live material is known, but I sus-
pect this may be a more arboreal species than most
Sinployea. Sinployea angularis (fig. 64d-f) is reduced
in size, has widely spaced radial ribbing, a narrow um-
bilicus, angulated periphery, and pale monochrome
coloration. It has perhaps the most modified shell of
any Sinployea. Possibly generic separation is war-
ranted, but without study of the soft parts, I prefer a
conservative treatment.
On Viti Levu, the monotypic genus Ba (p. 172) can
be derived from Sinployea irregularis by a drastic
change in coiling pattern, reduction in whorl count,
closure of the umbilicus, and decrease in number of
spiral apical cords. Anatomical changes are covered
under the description of that species. The other Viti
Levu species, Sinployea godeffroyana (fig. 69a-c), is
characterized by its small size, proportionately thick
body whorl, and extremely crowded radial ribbing
(mean ribs/mm. 28.4). It has not been dissected.
Undoubtedly there are additional species of
Sinployea in both Fiji and the Lau Archipelago. Use of
the following key can serve only as a general guide to
affinities when more collections become available.
KEY TO THE FIJI AND LAU Sinployea
1. Main islands of Fiji 2
Lau Archipelago 4
2. Ribbing irregular; apical cords more than 15; mean diameter
more than 2.7 mm Sinployea irregularis (Garrett, 1887)
Ribbing regular; apical cord average 10; mean diameter less
than 2.5 mm 3
3. Viti Levu; mean diameter about 2.1 mm.; ribs/mm, more than
20 Sinployea godeffroyana, new species
Taveuni and Kandavu; mean diameter about 2.3-2.4 mm.;
ribs/mm, less than 15 Sinployea princei (Liardet, 1876)
4. Mean diameter less than 3.0 mm.; mean D/U ratio less than
7 5
Mean diameter more than 3.1 mm.; mean D/U ratio more than
10 Sinployea adposita (Mousson, 1870)
5. Body whorl not angulated; mean diameter more than 2.3
mm 6
Body whorl angulated; mean diameter about 2.1 mm.;
Namuka Sinployea angularis, new species
6. Prominent major ribbing continuing to end of body whorl 7
Major radial ribbing widely spaced on spire, becoming reduced,
then absent on body whorl Sinployea recursa, new species
7. Mean rib count 100-150 8
Mean rib count 50-95 9
8. Mean D/U ratio less than 3.5; ribs/mm, more than 16; mean
body whorl width about 0.75 mm
Sinployea navutuensis, new species
Mean D/U ratio more than 4.0; ribs/mm, less than 15; mean
body whorl width about 0.90 mm
Sinployea lauensis, new species
9. Mean D/U ratio usually much more than 3.8; Central and
Northern Lau 10
Mean D/U ratio less than 3.6; Southern Lau
Sinployea inermis meridionalis, new subspecies
10. Mean rib count less than 60; mean D/U ratio about 5.50; mean
apical cords about 13; Lakemba
Sinployea inermis lakembana, new subspecies
Mean rib count more than 70; mean D/U ratio less than 4.50;
mean apical cords about 9; Mango and Vanua
Mbalavu Sinployea inermis inermis (Mousson, 1870)
Sinployea angularis, new species. Figure 64d-f.
Diagnosis. Shell very small, diameter 2.02-2.27 mm. (mean
2.11 mm.), with 3%-4 normally coiled whorls. Apex and spire mod-
erately and usually evenly elevated, sometimes rounded above, body
whorl descending a little more rapidly, spire protrusion almost Va
body whorl width, H/D ratio 0.512-0.615 (mean 0.574). Apical
sculpture of 13-16 (mean 14.6) narrow and rather inconspicuous
spiral cords. Postnuclear sculpture of low, rounded, widely spaced,
strongly protractively sinuated radial ribs, 36-42 (mean 38.0) on the
body whorl, whose interstices are 4-7 times their width. Ribs/mm.
5.28-6.16 (mean 5.67). Microsculpture of extremely fine radial rib-
lets, more than 10 between each pair of major ribs, crossed by equally
fine spiral riblets, with narrow, rather crowded secondary spiral
cording, becoming broader and less distinct on body whorl. All micro-
sculpture barely visible at 96 x magnification. Umbilicus narrow,
U-shaped, last whorl decoiling more rapidly, contained 5.21-8.71
times (mean 6.14) in the diameter, margins rounded. Sutures im-
pressed, whorls strongly rounded above, becoming strongly flattened
laterally above obtusely angulated periphery, evenly rounded on
FIG. 64. a-c, Sinployea recursa, new species. Station 27, Wangava, Lau, Fiji. Holotype. BPBM 166967; d-f, Sinployea angularis, new
species. Station 3, Namuka, Lau, Fiji. Holotype. BPBM 166616. Scale lines equal 1 mm. (MM).
148
SYSTEMATIC REVIEW
149
outer and lower margins. Aperture inclined about 25 from shell
axis.
Although both Sinployea irregularis from Viti
Levu (fig. 65e) and S. rudis from Rarotonga (fig. 47e)
give the appearance of having an angulated periphery
because of supraperipheral flattening, only S. an-
gularis (fig. 64e) of Polynesian-Fijian Charopidae has
the periphery itself actually angulated. Sinployea ad-
posita (fig. 69d-f) agrees in rib spacing, but is much,
much larger and has a very narrow umbilicus. Sin-
ployea inermis (fig. 66a-e) is much larger, with a less
protruded spire, rounded periphery, and usually much
more numerous and crowded ribbing.
Description. Shell very small, with 4 moderately tightly coiled
whorls. Apex and spire distinctly and evenly elevated, last part of
body whorl slightly deflected below periphery of penultimate, H/D
ratio 0.591. Embryonic whorls 1%, sculpture mainly eroded, with
traces of fine spiral cords remaining. Postnuclear sculpture of low,
rounded, protractively sinuated radial ribs, 36 on the body whorl,
whose interstices are 4-7 times their width. Microsculpture of ex-
tremely fine radial riblets, crossed by much finer, more crowded spi-
ral riblets and narrow, rather vague secondary spiral cords. Sutures
relatively shallow on body whorl, slightly indented. Whorls with
angulated periphery, somewhat flattened above and slightly flat-
tened below with gently rounded basal margin. Umbilicus narrow,
U-shaped, last whorl decoiling more rapidly, contained 5.50 times in
the diameter. Color leached from shell. Aperture ovate with angu-
lated periphery, inclined about 25 from the shell axis. Height of
holotype 1.28 mm., diameter 2.17 mm.
Holotype. Fiji: Lau Group, Namuka, Station 3,
500 ft. inland from Matandolo on north central part of
island at 65-75 ft. elevation. Collected dead between
rocks by H. S. Ladd on July 7, 1934. BPBM 166616.
Range. Namuka, Lau Group, Fiji.
Paratypes. Namuka: 500 ft. inland from Matan-
dolo (Station 3) on north central part of island at 65-75
ft. elevation (187 specimens, BPBM 166616, BPBM
166642, BPBM 166669).
Remarks. Only 14 (7.4%) of 188 Sinployea an-
gularis were whole adults. The remainder were
juveniles or badly broken specimens. Although an an-
gulated periphery is extremely common in the En-
dodontidae and many of the New Caledonian and
Austro-Zelandic endodontoid taxa, it is found other-
wise among Pacific Island Charopidae only in the
Marianas Himeroconcha quadrasi and H. fusca (fig.
105b, e). Hence the choice of angularis as specific name
for this species.
Its elevated spire and angulated periphery sepa-
rate it from any other Fijian species in appearance, but
the pattern of sculpture is a modification of the same
type seen in S. inermis. Sinployea angularis is much
smaller than examples of S. inermis, coming closest in
size to the Viti Levu S. godeffroyana. The latter has
very numerous and crowded radial ribbing, plus the
spire is much less protruded.
Possibly this is an arboreal species and should be
generically separated from Sinployea. Without dissec-
tion and ecological data, I prefer to retain it in
Sinployea. In size and shape it is the most drastically
altered species.
Sinployea recursa, new species. Figure 64a-c.
Diagnosis. Shell of average size, diameter 2.60-3.09 mm.
(mean 2.83 mm.), with 3 l /2-4'/8 normally coiled whorls. Apex and
spire moderately to strongly elevated, body whorl descending more
rapidly, spire protrusion more than Vi body whorl width, H/D ratio
0.562-0.634 (mean 0.587). Apical sculpture of 14-18 (mean 15. 9) fine
spiral cords, often partly to completely eroded. Postnuclear sculpture
of indistinct, low, broadly rounded, widely spaced radial ribs on the
upper spire that become greatly reduced to absent by early part of
body whorl. Vague traces of microreticulations on the upper spire
become replaced by fine and crowded secondary spiral cords on body
whorl. All microsculpture only visible under high magnification with
oblique lighting. Umbilicus narrow, V-shaped, slightly and regularly
decoiling, contained 5.75-7.90 times (mean 6.63) in the diameter,
margins rounded. Sutures impressed, whorls strongly rounded
above, compressed laterally above periphery, with slightly com-
pressed basal margin. Color light yellow-horn, with irregular red-
dish flammulations that become narrower on shell base.
In spire elevation and general appearance, Sin-
ployea recursa shows many similarities to S. angularis.
The latter (fig. 64d-f) is much smaller, has an angu-
lated periphery, and much more prominent radial rib-
bing. No other Polynesian or Micronesian Charopidae
has the ribbing so greatly reduced, although this is
frequent in New Caledonian, Australian, and New
Zealand taxa and occurs sporadically in the Polynesian
Endodontidae.
Description. Shell of average size, with slightly less than 3%
relatively loosely coiled whorls. Apex and spire evenly elevated, last
whorl descending a little more rapidly, H/D ratio 0.582. Embryonic
whorls 1!4, sculpture of about 15 fine spiral ribs, partially eroded.
Postnuclear whorls with a few widely spaced, low, inconspicuous
radial ribs on the upper spire, becoming reduced over most of the
body whorl but reappearing on last quarter. Microsculpture of fine
radial growth striae and very low, inconspicuous secondary spiral
cording. Sutures relatively shallow, whorls gently rounded above,
sloping flatly down to more strongly rounded periphery and slightly
flattened basal lip, contained 7.82 times in the diameter. Color light
yellowish white with vague irregular yellow-red or reddish brown
flammulations. Aperture ovate, slightly flattened basally, inclined
about 20 from the shell axis. Height of holotype 1.64 mm., diameter
2.83 mm.
Holotype. Fiji: Lau Group, Wangava, Station 27,
northeast end of island on an inland ridge, % mile from
shore, at 75 ft. elevation. Collected in soil under leaves
and between rocks by H. S. Ladd on July 22, 1934.
BPBM 166967.
Range. Wangava, Lau Group, Fiji.
Paratypes. Wangava: northeast end of island
(Station 27), % mile from shore on ridge at 75 ft. eleva-
tion (107 specimens, BPBM 166967, BPBM 166969,
BPBM 166996).
Remarks. The similarity of Sinployea recursa to
the Austro-Zelandic flammulinid taxa with reduced
ribbing is striking, but the probabilities are that this is
convergent. Sinployea angularis is intermediate in
sculpture between S. recursa and the Fijian charopids
with typical sculpture, so that I have no hesitation in
considering it to be a local development. No soft parts
were available for dissection.
No other Charopidae covered in this monograph
has such sculptural reduction. Although many species
have the ribbing extremely irregular, such Fijian
150
SOLEM: ENDODONTOID LAND SNAILS
shells as S. irregularis (fig. 65d-f) and Ba humbugi
(fig. 74a-c) being good examples, the almost complete
loss of ribbing does not occur. This condition is more
common in the Endodontidae, where a number of En-
dodonta, several Nesodiscus, one Nesophila, and one
Libera have undergone such reduction. In all of these
cases, the shells are quite large, and sculptural reduc-
tion is associated with size increase. The name recursa
refers to the reduced shell sculpture on the lower
whorls in particular, indicating a return to simplified
shell structure.
I suspect it may be an arboreal or semiarboreal
species.
Sinployea princei (Liardet, 1876). Figure 65a-c.
Helix princei Liardet, 1876, Proc. Zool. Soc. London, 1876, p. 100,
pi. 5, fig. 4, a Taveuni Island, Fiji Islands.
Patula princei (Liardet), Garrett, 1887, Proc. Zool. Soc. London,
1887, p. 179.
Helix (Patula) princei Liardet, Tryon, 1887, Man. Conchol., (2) 3, p.
27, pi. 5, fig. 83a.
Endodonta (Charopa) princei (Liardet), Pilsbry, 1893, Man. Con-
chol., (2)9, p. 35.
Champa princei (Liardet), Gude, 1913, Proc. Malacol. Soc. London,
10, (5), p. 330; Germain, 1932, Ann. Inst. Ocean., 12 (2), p. 45.
Diagnosis. Shell small, diameter 2.35-2.68 mm. (mean 2.43
mm.), with 3Vz-4Vn rather tightly coiled whorls. Apex and spire mod-
erately and evenly elevated, last whorl descending more rapidly,
spire protrusion more than Ve body whorl width, H/D ratio 0.524-
0.586 (mean 0.551). Apical sculpture of 8-10 (mean 9.0) fine and
relatively low spiral cords. Postnuclear sculpture of high, prominent,
thin, protractively sinuated radial ribs, 80-102 (mean 91.1) on the
body whorl, whose interstices are l'/2-3 times their width. Ribs/mm.
11.85-13.80 (mean 12.06). Microsculpture of fine radial riblets, 3-6
between each pair of major ribs, crossed by extremely fine and
crowded spiral riblets that are barely visible under 96x magnifica-
tion. Secondary spiral cording occasionally visible near periphery
and shell base. Umbilicus open, U-shaped, last whorl decoiling more
rapidly, contained 3.74-4.83 times (mean 4.28) in the diameter,
margins rounded. Whorls very strongly rounded, aperture circular,
sometimes slightly flattened laterally above periphery, inclined
about 20 from shell axis. Color reddish horn, no flammulations.
Sinployea princei is very similar to S. inermis in-
ermis (fig. 66a-c), differing primarily in its much more
crowded radial ribbing, greatly reduced secondary spi-
ral cording, and slightly smaller size. Sinployea lauen-
sis (fig. 68a-c) is larger, flatter, and has a distinctly
less protruding spire. Other Fijian species differ
greatly in rib spacing and count. Sinployea ellicensis
(fig. 62a-f) has a flammulated color pattern, much
more prominent secondary spiral cording, and aver-
ages three-eighths of a whorl less.
Description. Shell small, with 3% moderately tightly coiled
whorls. Apex and spire slightly and evenly elevated, body whorl
descending more rapidly, H/D ratio 0.545. Apical whorls 1%,
sculpture of 8 narrow, widely spaced spiral ribs with faint traces of
weaker radial riblets on last eighth whorl. Postnuclear whorls with
narrow, lamellar, somewhat crowded, slightly protractively sinuated
radial ribs, 102 on the body whorl, whose interstices are 2-4 times
their width. Microsculpture of weak radial and finer spiral riblets
with a secondary sculpture of weak, widely spaced spiral cords visi-
ble only on periphery and base of body whorl. Sutures deep, whorls
strongly rounded above, evenly rounded on outer and basal margin.
Color light reddish yellow-horn without darker markings. Umbilicus
narrow, U-shaped, regularly decoiling, contained 4.08 times in the
diameter. Aperture subcircular with evenly rounded outer margin,
inclined about 15 from the shell axis. Height of lectotype 1.28 mm.,
diameter 2.36 mm.
Lectotype. Fiji: Taveuni Island. Collected by
Liardet. BMNH 76.9.9.2.
Range. Taveuni and Kandavu, Fiji Islands.
Paratypes.BMNH 76.9.9.2.
Material. Fiji (2 specimens, Brussels): Taveuni (6
specimens, Zurich); Kandavu (4 specimens, FMNH
48997, SMF 165749).
Remarks. Specimens from Kandavu, originating
from the Rolle collection, are separable (table XXX)
from the Taveuni specimens only by their narrower
umbilicus. They are considered to be S. princei, but
were not included in figuring mean size because I am
uncertain that the locality is accurate.
No material of S. princei has been collected in this
century.
Sinployea inermis (Mousson, 1870).
Several populations from islands in the Lau Group
are considered to be conspecific. They differ from the
Taveuni and Kandavu S. princei (fig. 65a-c) in their
higher spire protrusion and much less crowded radial
ribbing (fig. 66a-e). Lau Archipelago species of similar
size, S. lauensis (fig. 68a-c) and S. navutuensis (fig.
68d-f), plus the Viti Levu S. godeffroyana (fig. 69a-c),
all have much more numerous and crowded radial rib-
bing plus moderately to greatly different proportions.
Extralimital species such as the Tongan S. vicaria (fig.
58a-c) and Ellice Island S. ellicensis (fig. 62a-c) differ
in details of sculptural character and spacing, but ob-
viously are closely related.
Variation among populations on the same island
and from different islands is relatively extensive (ta-
bles XXXII- XXX VI) and somewhat difficult to inter-
pret. Available material was collected in the 1850s,
1860s, in 1934, 1938, 1973, and 1977. Since the Lau
Group has unpredictable periods of drought, the possi-
bility of ecophenotypic dwarfing must be considered in
accessing the mean differences. Most samples fit rela-
tively well into one of three geographic variation pat-
terns that are considered to represent subspecies.
Specimens from the northern group of Vanua Mbalavu,
Mango, and Kimbombo Islands are nearest to S. prin-
cei, having more ribs and intermediate umbilical width
(fig. 66a-c). They average about nine apical cords.
Samples from the large, centrally located island of
Lakemba vary in umbilical width but average a nar-
row umbilicus, 11.6 apical cords, and have very widely
spaced and large radial ribs (fig. 66e). Those from the
more southerly Lau Group, Aiwa, Yangasa Levu, and
Navutu-i-Loma tend to have a slightly wider um-
bilicus, average 13.6 apical cords, have a little less
crowded ribbing than those from the northern area,
and are easily separable from the Lakemba popula-
tions (fig. 66d). Frequency distributions of the D/U
ratio, ribs/mm., and ribs (tables XXXII- XXXIII) illus-
trate the extent of overlap for any single character. Rib
a
FIG. 65. a-c, Sinployea princei (Liardet). Taveuni, Fiji. Probable paratype. Zoologisches Institut der Universitat Zurich; d-f, Sinployea
irregularis (Garrett). Viti Levu, Fiji. Lectotype. BPBM 7982. Scale lines equal 1 mm. (MM).
151
TABLE XXXI. - LOCAL VARIATION IN FIJIAN, MELANESIAN AND MICRONESIAN SINPLOYEA.
NUMBER OF
SPECIMENS
RIBS/MM.
DIAMETER
H/D RATIO
Sta. 3
BPBM 166642
Sta. 2
BPBM 166616
1.230.025 2.09*0.024 0.58810.0121
(1.10-1.28) (2.01-2.17) (0.540-0.615)
1.1810.028 2.1010.031 0.56110.0090
(1.07-1.28) (2.02-2.25) (0.512-0.591)
Sta. 27
BPBM 166967
lauensis
12
4
5
3
Sta. 30
BPBM 167139
Sta. 43
BPBM 167234
Sta. 28, 29
BPBM 167020, -55
navutuensis
Sta. 29 4
ex BPBM 167020,
ex BPBM 167055
kusaieana
Sta. 44 11
BPBM 155874-5, -7, -9
descendens
Karlei 24
ZMB
Matong 7
ZMB
100.412.23
(95-105)
119
142
11.5910.287
(10.96-12.62)
12.51
11.64
1.6610.027 2.8110.045 0.58910.0066
(1.52-1.85) (2.62-3.11) (0.562-0.634)
1.3810.065 2.6810.112 0.51510.0110
(1.19-1.49) (2.42-2.95) (0.493-0.544)
1.4810.076 2.7910.078
0.53110.0143
(1.39-1.79)
(2.65-3.08)
(0.494-0.581)
1.6110.145 2.9610.141 0.54210.0242
(1.32-1.79) (2.68-3.15) (0.494-0.568)
1.3210.73 2.6610.048 0.49810.0220
(1.16-1.49) (2.58-2.78) (0.449-0.556)
90.411.97 11.1910.287 1.3110.027 2.5810.020 0.50610.0074
(84-103) (9.96-13.17) (1.18-1.48) (2.50-2.73) (0.481-0.542)
91.711.55 13.0710.232 1.1610.009 2.2510.013 0.51510.0041
(77-103) (11.16-14.71) (1.05-1.25) (2.14-2.40) (0.470-0.548)
91.912.62 12.6810.313 1.2210.017 2.3210.037 0.52810.0062
(80-99) (11.33-13.73) (1.15-1.27) (2.15-2.43) (0.504-0.550)
novopomme rana
Karlei
ZMB
12 93.813.29 12.7110.499 1.2110.044 2.3310.059 0.52010.0089
(72-l'04) (9.61-15.10) (1.05-1.61) (2.11-2.85) (0.467-0.567)
WHORLS
UMBILICUS
D/U RATIO
APICAL CORDS
Sta. 2
3 7/8+(3 3/4-4)
3 7/8-O 3/4-4)
0.3210.531
(0.23-0.36)
6.7110.415 15
(5.81-8.71)
0.3810.007 5.5710.099
(0.35-0.39) (5.21-6.05)
recursa
Sta. 27
lauensis
Sta. 30
Sta. 43
Sta. 28
navutuensis
Sta. 29
kusaieana
Sta. 44
descendens
Karlei
Ma t ong
3 7/8-(3 1/2-4 1/8) 0.4210.015 6.7010.211 16.511.50
(0.33-0.49) (5.78-7.90) (15-18)
3 5/8+(3 1/2-3 7/8) 0.6110.039 4.4010.120
(0.53-0.70) (4.15-4.65)
3 7/8(3 3/4-4 1/8) 0.6810.030 4.1010.085
(0.63-0.79) (3.88-4.32)
4 1/8(3 7/8-4 1/4) 0.6910.038 4.2610.150
(0.63-0.76) (4.00-4.52)
4 1/8(4-4 3/8) 0.7810.016 3.3810.055 10. Oil. 00
(0.76-0.82) (3.25-3.52) (9-11)
4-(3 3/4-4 1/8) 0.7410.009 3.5110.038 9.210.20
(0.69-0.79) (3.21-3.67) (8-10)
4 l/8+(4-4 1/4) 0.8310.011 2.7310.029 10.710.30
(0.76-0.95) (2.43-2.94) (9-14)
4 3/8-(4 1/8-4 1/2) 0.8510.028 2.7310.046 10.510.29
(0.74-0.95) (2.55-2.91) (10-11)
3 3/4(3 1/2-4 1/8) 0.6710.028 3.4910.076 10.710.36
(0.54-0.92) (3.09-3.96) (10-12)
152
SYSTEMATIC REVIEW 153
TABLE XXXII. - D/U RATIO FREQUENCY IN SINPLOYEA INERMIS SUBSPECIES. TABLE XXXIV. - RIB FREQUENCY IN SINPLOYEA INERMIS SUBSPECIES.
D/U Ratio
inermis
3.00-3
.39
3.40-3
.79
7
3.80-4
.19
17
4.20-4
.59
17
4.60-4
.99
6
5.00-5
.39
5.40-5
.79
5.80-6
.19
6.20-6
.59
6.60-6
.99
7.00-7
.39
meridionalis
7
17
2
S. inermis
lakembana
Ribs
35-41
42-48
49-55
56-62
63-70
71-77
78-84
85-91
92-98
99-105
S . inermis
inermis
meridionalis
differences are somewhat obscured by the strong ten-
dency for gerontic crowding and irregularity of rib
growth in most adult specimens. The differences are
much clearer on the penultimate and first part of the
body whorl, but convenient quantification of data from
this section of the shell was not achieved. Most speci-
mens are readily assignable to one of the three groups.
Hence, recognition of three subspecies is proposed:
Sinployea inermis inermis from Vanua Mbalavu, Kim-
bombo, and Mango; S. i. lakembana from Lakemba;
and S. i. meridionalis from Aiwa, Navutu-i-Loma, and
Yangasa Levu.
Larger collections from more islands may result in
altering this classification. The apical cording counts
and the size differences in ribbing are substantial. Pos-
TABLE XXXIII. - RIBS/MM. FREQUENCY IN SINPLOYEA INERMIS
SUBSPECIES.
S . inermis
Ribs/mm.
inermis
4.50-4.99
5.00-5.49
5.50-5.99
6.00-6.49
6.50-6.99
7.00-7.49
7.50-7.99
1
8.00-8.49
1
8.50-8.99
5
9.00-9.49
2
9.50-9.99
5
10.00-10.49
2
10.50-10.99
4
11.00-11.49
2
11.50-11.99
1
S. inermis
meridionalis
lakembana
sibly specific level differentiation is involved, but the
presence of samples not clearly referable to a sub-
species and the amount of overlap (tables XXXII-
XXXVI) suggest conservative treatment.
The specimens collected from "Vatou Island"
(= Vanua Vatu) during the surveys of the Herald in the
mid-1850s are still preserved in the British Museum
(Natural History) (BMNH 59.1.14.63). Only one fully
adult shell is in good condition, but in general appear-
ance the specimens are intermediate between sub-
species inermis and subspecies lakembana. They have
the relatively widely spaced ribbing that is char-
acteristic of meridionalis and lakembana, although
somewhat more crowded than typical examples of
either subspecies. The umbilicus varies widely in
width from the narrow type of lakembana to the widely
opened umbilicus of meridionalis.
Single shells from two islands lying within the
range of S. i. meridionalis could not be assigned to a
subspecies with assurance. Their measurements are
given in Table XXXVI. The Ongea example (BPBM
161228) has a narrower umbilicus than does the nomi-
nate race or any specimens of S. i. meridionalis but
disagrees in sculpture and apical cording with S. i.
lakembana. An adult shell from Tavu-na-Sithi (BPBM
78805) could be referred to either typical inermis or
meridionalis. It has the umbilical size of inermis, but
the ribbing is closer to that of meridionalis. Collection
of adequate samples is needed in order to establish the
position of these populations.
Derivation of S. adposita (fig. 69d-f) from S. iner-
mis requires size increase, greater protrusion of the
spire and its correlated narrowing of the umbilicus,
plus greater spacing of the ribs. Similarly, increase in
body whorl descension and development of ribbing ir-
regularity would be sufficient to derive S. irregularis
(fig. 65d-f) from either S. inermis or S. princei.
The penis of Sinployea inermis is slightly more
than half the size of the penis inS. adposita and differs
radically in shape and internal proportions. The vergic
154
SOLEM: ENDODONTOID LAND SNAILS
TABLE XXXV. - LOCAL VARIATION IN SINPLOYEA INERMIS, I.
NUMBER OF
SPECIMENS
inermis inermis
Vanua Mbalavu
Sta. 70, 72(1938)
Sta. 69(1938)
BPBM 179396
Ex Fulton
BPBM 167433
Mango
Sta. 89(1938)
BPBM 179941
Sta. 91(1938)
BPBM 180025-6
Kimbombo
Bryan! (1924)
BPBM 79103
Inermis meridionalis
Yangasa Levu
Sta. 30(1934)
BPBM 167138
Aiwa
Sta. 40(1934)
BPBM 167209
6
10
14
10
75.35.37
(65-83)
76.31.33
(75-79)
78.0+8.61
(65-102)
78.02.01
(76-80)
91.54.72
(77-104)
79
72
93.011.03
(82-104)
HEIGHT
1.28+0.020
(1.23-1.36)
DIAMETER
2.28+0.013
(2.25-2.32)
1.360.032 2.39+0.029
(1.19-1.56) (2.28-2.55)
1.48+0.058 2.52+0.067
(1.36-1.66) (2.38-2.78)
1.400.065 2.56+0.064
(1.19-1.62) (2.32-2.78)
1.67+0.032 2.950.033
(1.46-1.85) (2.75-3.08)
1.45+0.067 2.640.140
(1.32-1.59) (2.28-2.88)
H/D RATIO
0.5620.0061
(0.544-0.586)
0.5700.0094
(0.514-0.627)
0.5850.0134
(0.547-0.627)
0.546+0.0127
(0.506-0.583)
0.564+0.0075
(0.530-0.629)
0.5650.0065
(0.558-0.580)
1.81+0.037 3.040.029 0.5950.0149
(1.59-1.95) (2.95-3.25) (0.505-0.655)
1.660.027 2.960.109 0.5620.0205
(1.59-1.72) (2.72-3.21) (0.521-0.610)
i.iner.
WHORLS
Mbalavu 3 5/8+(3 1/2-3 7/8)
Sta. 70,72
Sta. 69 3 7/8(3 5/8-4 1/4)
Ex Fulton 4-(3 7/8-4 1/8)
Mango
Sta. 89
Sta. 91
Kimbombo
Bryan!
i.merid.
Yangasa
Sta. 30
4 l/8+(3 7/8-4 1/2)
4 l/4-(4-4 5/8)
3 7/8-(3 1/2-4 1/8)
4 3/8+(4 1/4-4 5/8)
Aiwa
Sta. 40 4 l/4-(4-4 1/4)
UMBILICUS
D/U RATIO
APICAL
CORDS
0.53+0.013 4.270.094 8.00+0.71
(0.49-0.59) (3.89-4.60) (7-10)
0.5810.016 4.0910.098 9.3810.26
(0.49-0.66) (3.50-4.60) (8-10)
0.570.026 4.45+0.112 8.331.20
(0.53-0.66) (4.17-4.69) (6-10)
0.65+0.034 3.940.139 9
(0.53-0.72) (3.55-4.38)
0.70+0.017
(0.63-0.82)
0.63+0.087
(0.49-0.79)
4.20+0.081 9.57+0.37
(3.68-4.65) (8-11)
4.15+0.260 9
(3.58-4.60)
0.8710.025 3.500.092 12.0010.58
(0.72-0.99) (3.27-4.14) (11-13)
0.810.044 3.57+0.013
(0.69-0.89) (3.54-3.60)
papilla in the former is about a third of the size of the
latter, and the muscular collar is larger both actually
and proportionately. Sinployea adposita has the stim-
ulatory pad smaller and shorter than in S. inermis. The
latter has a much broader penis that is wider medially
than apically and then tapers abruptly to atrium.
Sinployea adposita has the penis enlarged apically and
medially, then it tapers gradually to atrium.
Sinployea inermis inermis (Mousson, 1870). Figures
66a-c, 67e.
Patula (Patula) inermis Mousson, 1870, J. de. Conchyl., 18, pp.
118-119, pi. 7, fig. 7 Lomma-Lomma on Vanua-Balavo, Lau
Group, Fiji Islands.
Helix inermis (Mousson), PfeifTer, 1876, Monog. helic. viv., 7, p.
167.
Patula inermis Mousson, Garrett, 1887, Proc. Zool. Soc. London,
1887, p. 178.
SYSTEMATIC REVIEW
TABLE XXXVI. - LOCAL VARIATION IN SINPLOYEA INERMIS, II.
155
NUMBER OF
SPECIMENS
inermis meridionalis
Navutu-I-Loma 12
Sta. 28-9(1934)
BPBM 167019,
-55-6
inermis lakembana
Transitional Sets
Vatou 5
BMNH 59.1.14.
63
Ongea
Tavua-na-
Sithi
RIBS
68.03.16
(53-83)
HEIGHT
1.480.39
(1.32-1.69)
DIAMETER
2.72+0.047
(2.52-3.05)
Lakemba
Sta. 42(1934)
BPBM 167222
4
54.
814.61
(42-62)
1.
48+0
(1
.083
.32-1
.66)
2,
.5810
(2
.111
.38-2.
88)
Sta. 97(1938)
BPBM 180101
10
49.
5+1.71
(38-59)
1.
3610
(1
.025
.19-1.
.49)
2,
,2910
(2
.028
.15-2.
38)
65.83.40
(59-70)
83
75
1.3310.093
(1.13-1.61)
1.41
1.58
2.420.100
(2.19-2.73)
2.66
2.70
H/D RATIO
0.544+0.0087
(0.500-0.595)
0.5740.0195
(0.552-0.633)
0.5940.0085
(0.545-0.634)
0.5480.0177
(0.518-0.616)
0.531
0.586
i.iner.
WHORLS
Navutu
Sta. 28-29 4 l/8+(3 7/8-4 1/2)
i . lakem.
Lakemba
Sta. 42 4 l/8+(3 7/8-4 1/2)
Sta. 97
4-(3 3/4-4 1/8)
Transitional
Vatou 4(3 7/8-4 1/4)
Ongea 3 3/4
Tavua. 4 1/4 -
UMBILICUS
D/U RATIO
APICAL
RIBS
0.78+0.021 3.50+0.055 11.430.68
(0.69-0.92) (3.07-3.81) (9-14)
0.620.049 4.2110.251
(0.53-0.76) (3.78-4.93)
0.3710.015 6.24+0.206 13.57+0.72
(0.33-0.49) (4.73-7.10) (11-16)
0.57+0.026 4.30+0.375
(0.49-0.66) (3.32-5.53)
0.56
0.66
4.77
4.10
Helix (Patula) inermis (Mousson), Tryon, 1887, Man. Conchol, (2)
3, p. 41, pi. 8, figs. 60-61.
Endodonta (Champa) inermis (Mousson), Pilsbry, 1893, Man. Con-
chol. (2) 9, p. 35.
Charopa inermis (Mousson), Gude, 1913, Proc. Malacol. Soc. Lon-
don, 10 (5), p. 330 Vatou (Herald Expedition!); Germain, 1932,
Ann. Inst. Ocean., 12 (2), p. 45.
Diagnosis. Shell relatively small, diameter 2.24-3.06 mm.
(mean 2.60 mm.), with 3 l /2-4% normally coiled whorls. Apex and
spire usually moderately and evenly elevated, last whorl descending
moderately to much more rapidly, spire protrusion about '/fe body
whorl width, H/D ratio 0.500-0.629 (mean 0.562). Apical whorls with
6-11 (mean 8.96) prominent spiral cords. Postnuclear whorls with
high, prominent, narrow, strongly protractively sinuated radial ribs,
65-104 (mean 80.0) on the body whorl, whose interstices are 3-6
times their width. Ribs/mm. 8.39-11.75 (mean 9.82). Microsculpture
of very fine radial riblets, 6-12 between each pair of major ribs,
crossed by slightly finer and more crowded spiral riblets and a few
widely spaced secondary spiral cords. Umbilicus open, U-shaped, last
whorl decoiling much more rapidly, contained 3.47-4.69 times (mean
4.17) in the diameter, margins rounded. Sutures and whorl contours
typical, color reddish yellow-horn without separate flammulations.
Sinployea inermis inermis (fig. 66c) has a narrower
umbilicus and more crowded major radial ribbing than
S. i. meridionalis (fig. 66d); S. i. lakembana (fig. 66e)
has much coarser and more widely spaced major radial
ribs. Other Fijian species of similar size, S. princei, S.
lauensis, and S. navutuensis, differ in proportions
(table XXX) and/or ribbing. The Tongan S. uicaria (fig.
58a-c) has a higher spire and much more crowded sec-
a
FIG. 66. a-c, Sinployea inermis inermis (Mousson). Station 72, Lomaloma, Vanua Mbalavu, Lau, Fiji. BPBM 179460; d, Sinployea inermis
meridionalis, new subspecies. Station 30, Yangasa Levu, Lau, Fiji. Holotype. BPBM 167138; e, Sinployea inermis lakembana, new subspecies.
Station 42, Lakemba, Lau, Fiji. Holotype. BPBM 167222. Scale lines equal 1 mm. (a-c, SH: d-e, MM).
156
SYSTEMATIC REVIEW
157
ondary spiral cording in addition to the much more
crowded radial ribbing.
Description. Shell relatively small, with slightly less than 4'/8
normally coiled whorls. Apex barely emergent, upper spire flat, body
whorl descending moderately sharply, H/D ratio 0.515. Apical whorls
1%, sculpture partially eroded with traces of 7 fine, rather widely
spaced spiral ribs remaining. Postnuclear whorls with prominent,
rounded, protractively sinuated radial ribs, 69 on the body whorl,
whose interstices are 2-4 times their width. Microsculpture of very
fine radial riblets, 4-6 between each pair of major ribs, crossed by
even finer, more crowded spiral riblets with a secondary sculpture of
moderately prominent, widely spaced spiral cords developed. Sutures
deep, whorls strongly rounded above, moderately flattened laterally
above periphery and on basal margin. Umbilicus open, U-shaped,
last whorl decoiling more rapidly, contained 3.47 times in the diame-
ter, rounded margins. Color light reddish brown. Aperture ovate, lip
slightly deflected above, flattened laterally above periphery and on
basal margin, inclined about 20 from the shell axis. Height of lec-
totype 1.38 mm., diameter 2.58 mm.
Lectotype. Fiji: Lau Group, Vanua Mbalavu. Col-
lected by Dr. Graeffe. JDCP 3214.
Range. Vanua Mbalavu, Mango, Kimbombo,
Tuvutha, Lau Group, Fiji.
Paratypes. Zurich, FMNH 116998.
Material. Vanua Mbalavu (6 specimens, BPBM
167433): Lomaloma (9 specimens, Zurich, J. de Con-
chyl., FMNH 116998); north end Lomaloma (Station
69) 10-12 ft. inland at 3 ft. elevation (28 specimens,
BPBM 179396, BPBM 179398, BPBM 179399); back of
Lomaloma (Station 67) l /2-% mile inland, 250-700 ft.
elevation (8 specimens, BPBM 179379); south end
Lomaloma (Station 72) 15 ft. inland at 3 ft. elevation
(14 specimens, BPBM 179460, BPBM 179461); south
end Unione Village (Station 70) 10 ft. inland at 3 ft.
elevation (2 specimens, BPBM 179426); between Val-
ika and Mosomo Bay (Station 78) % mile inland at
200-250 ft. elevation (3 specimens, BPBM 179584,
BPBM 179585); Station F-8, 300 ft. elevation, P/2 miles
inland, northwest area (6 specimens, FMNH 168161).
Mango Island: Marona (Station 92) 100 yd. inland at 10
ft. elevation (2 specimens, BPBM 180047); south-
southwest from Marona (Stations 89, 90) about Vz-%
mile inland at 200-400 ft. elevation (15 specimens,
BPBM 179941, BPBM 179975, BPBM 179988); %-l
mile southwest of Marona (Station 91) 1-50 yd. inland,
at 5-30 ft. elevation (82 specimens, BPBM 180025-9).
Kimbombo: East Islet, 100 yd. inland at 50-150 ft. ele-
vation (28 specimens, BPBM 79079, BPBM 79103).
Tuvutha: west coast (Station F-7) at 100 ft. elevation (5
specimens, FMNH 193768).
Remarks. The specimens of Sinployea inermis
inermis from Vanua Mbalavu, Mango, and Kimbombo
agree in sculpture and umbilical proportion, although
exhibiting slight differences in size (table XXXV).
Shells from Vanua Mbalavu are small, whereas those
from Kimbombo and Mango are slightly to distinctly
larger. On Mango, two populations were sampled.
Shells from Station 89 (Vz-% mile south-southwest of
Marona at 350-400 ft. elevation) were dead. They are
noticeably smaller and with fewer radial ribs than
those from Station 91 (%-l mile southwest of Marona
at 5-30 ft. elevation), which were collected alive. The
latter sample contained a higher proportion of gerontic
individuals, whereas the Station 89 material was
barely adult. The Mango specimens are larger than
those from Vanua Mbalavu, but no hesitation is felt in
referring them to the same subspecies.
Material from both Mango (BPBM 180025) and
Vanua Mbalavu (BPBM 179396) was dissected. Only
differences are noted below.
Description of soft parts. Pallial region typical, arms of kidney
subequal, rectal slightly longer, maximum length 1.7 mm., slight
strip of lung roof visible between ureter arms.
Genitalia (fig. 67e) typical, free oviduct slightly more slender
than usual. Penis (P) about 0.8-1.0 mm. long, greatly swollen me-
dially. Internally with very tiny vergic papilla surrounded by a
heavy muscular collar. Stimulatory pad very large, walls below pad
with obscure pilasters.
Sinployea inermis meridionalis, new subspecies.
Figure 66d.
Diagnosis. Shell of average size, diameter 2.50-3.22 mm.
(mean 2.88 mm.), with 3%-4% rather tightly coiled whorls. Apex and
spire moderately and evenly elevated, body whorl descending mod-
erately to much more rapidly, spire protrusion about l k body whorl
width, H/D ratio 0.505-0.655 (mean 0.566). Apical sculpture of 9-14
(mean 11.6) prominent, but narrow spiral cords. Postnuclear
sculpture of narrow, prominent, protractively sinuated, rather
widely spaced radial ribs, 53-104 (mean 72.2) on the body whorl,
whose interstices are 3-7 times their width. Ribs/mm. 6.75-11.70
(mean 8.37). Microsculpture of very fine radial riblets, 8-12 between
each pair of major ribs, crossed by barely visible spiral riblets, with a
weak secondary sculpture of fine and widely spaced spiral cords that
varies in frequency and prominence. Umbilicus broadly U-shaped,
last whorl decoiling much more rapidly, contained 3.07-3.81 times
(mean 3.51) in the diameter, margins rounded. Whorls strongly flat-
tened laterally above periphery, aperture subcircular, inclined about
20 from shell axis.
Sinployea inermis meridionalis (fig. 66d) has a
wider umbilicus and more widely spaced major radial
ribbing than the nominate race; S. i. lakembana (fig.
66e) is smaller, has a narrower umbilicus, and much
more widely spaced radial ribbing. Sinployea vicaria
paucicosta (fig. 58d) from Tonga is smaller, has the
spire protruding further, and has a narrower um-
bilicus.
Description. Shell slightly smaller than average, with 4'/4 rela-
tively tightly coiled whorls. Apex barely elevated, lower whorls de-
scending slowly, body whorl more rapidly, H/D ratio 0.565. Em-
bryonic whorls l'/2, sculpture of 11 relatively fine spiral ribs with an
intrusion of faint radial ribbing on the last quarter whorl. Remain-
ing whorls with rounded, prominent, protractively sinuated radial
ribs, 67 on the body whorl, whose interstices are about 3-6 times
their width. Microsculpture of very fine radial riblets, 8-12 between
each pair of major ribs, with exceedingly fine, inconspicuous spiral
riblets and a secondary sculpture of low, broadly rounded, relatively
widely spaced spiral cords. Sutures deeply impressed, whorls
strongly rounded above, flattened laterally above peripheral margin,
slightly compressed basally. Umbilicus broadly open, last whorl de-
coiling more rapidly, contained 3.29 times in the diameter. Color
light yellow-reddish brown without darker flammulations. Aperture
circular, strongly flattened laterally above periphery, with sharply
rounded umbilical basal margin, inclined about 10 from the shell
axis. Height of holotype 1.71 mm., diameter 3.03 mm.
158
SOLEM: ENDODONTOID LAND SNAILS
DG
FIG. 67. Anatomy of Fijian, Caroline Island, and New Hebridean Sinployea: a-d, Sinployea irregularis (Garrett). Station 118, Nandarivatu,
Viti Levu, Fiji. BPBM 178897. a, genitalia. 6, detail of talon-carrefour region, c, partial view of penis interior, d, apical part of penis interior; e,
genitalia of Sinployea inermis inermis (Mousson). Station 91, Marona, Mango Island, Lau, Fiji. BPBM 180025; f, genitalia of Sinployea inermis
lakembana, new subspecies. Station 97, Toumba, Lakemba Island, Lau, Fiji. BPBM 180101; g, penis and epiphallus of Sinployea adposita
(Mousson). Station 97, Toumba, Lakemba Island, Lau, Fiji. BPBM 180099; h, genitalia of Sinployea kusaieana, new species. Station 44, Lele
Islet, Kusaie, Caroline Islands. BPBM 155874; i-j, Sinployea euryomphala (Solem). Espiritu Santo, New Hebrides. FMNH 109493. (a-c, e-f, h,
MO; d, MM; g, JC; i-j, PS).
Holotype. Fiji: Lau Group, Yangasa Levu, Sta-
tion 30, hillside on south end of island, 150 ft. inland at
40 ft. elevation. Collected by H. S. Ladd on July 26,
1934. BPBM 167138.
Range. Yangasa Levu, Navutu-i-Loma, and
Aiwa, Lau Group, Fiji.
Paratypes. Yangasa Levu: hillside at south end
of island (Station 30), 150 ft. inland at 40 ft. elevation
(96 specimens, BPBM 167138). Aiwa: Vataniyambia
on southwest quarter (Station 40) of island, about 50
yd. inland at 25 ft. elevation (20 specimens, BPBM
167209). Navutu-i-Loma: ridge on northeast quarter of
SYSTEMATIC REVIEW
159
island (Stations 28, 29), about 150 yd. inland at 100 ft.
elevation (52 specimens, BPBM 167019, BPBM
167056, BPBM ex 167055).
Remarks. The shells of Sinployea inermis mer-
idionalis from Navutu-i-Loma are distinctly smaller
and more depressed in H/D ratio than are those from
Aiwa or Yangasa Levu (tables XXXV-XXXVI). Shells
from Aiwa have a much higher rib count. Compara-
tively few of the examples were adult or whole enough
to measure, so that the significance of this data is un-
certain. From Navutu-i-Loma there were 23.1% adults;
from Aiwa 20%; and from Yangasa Levu only 10.3%. I
would expect that the size difference is real, but more
complete collections are required.
Sinployea inermis lakembana, new subspecies. Fig-
ures 66e, 67f.
Diagnosis. Shell small, diameter 2.14-2.86 mm. (mean 2.36
mm.), with 3%-4'/2 rather tightly coiled whorls. Apex and spire mod-
erately to strongly and evenly elevated, last whorl descending much
more rapidly, spire protrusion more than l k body whorl width, H/D
ratio 0.508-0.634 (mean 0.583). Apical sculpture of 11-16 (mean
13.6) prominent spiral cords. Postnuclear sculpture of high, very
prominent, sharply defined, protractively sinuated radial ribs, 38-64
(mean 51.3) on the body whorl, whose interstices are 4-6 times their
width on the spire and first part of body whorl, becoming crowded
and irregular on gerontic growth sections. Ribs/mm. 4.67-8.20
(mean 7.01). Microsculpture of extremely fine radial riblets, more
than 10 between each pair of major ribs, but usually too fine for
accurate counting at 96 x magnification, barely visible microspirals,
and more prominent, rather widely spaced secondary spiral cording.
Umbilicus variable in size, usually narrow, U-shaped, last whorl
decoiling slightly to much more rapidly, contained 3.07-7.10 times
(mean 5.54) in the diameter, margins strongly rounded. Sutures very
deep, whorls shouldered above, compressed laterally above periphery
and on basal margin, aperture inclined about 30 from shell axis.
Sinployea inermis lakembana has much stronger
and more widely spaced major radial ribbing, plus a
usually more strongly deflected body whorl and nar-
rower umbilicus than either of the other races of S.
inermis (fig. 66a-d). No other Fijian species has such
strong ribbing that is so widely spaced.
Description. Shell relatively small, with 4'/2 tightly coiled
whorls. Apex very slightly elevated, lower spire and body whorl de-
scending more rapidly, H/D ratio 0.552. Apical whorls l'/2, sculpture
of 12 prominent spiral cords, surface partially eroded. Postnuclear
whorls with prominent, relatively narrow, quite widely spaced radial
ribs, 42 on the body whorl, whose interstices are 3-6 times their
width. Microsculpture of extremely fine radial and even finer, more
crowded spiral riblets, crossed by low, rounded, relatively closely
spaced spiral cords. Sutures deeply impressed, whorls strongly
rounded, only slightly flattened laterally above periphery and on
basal margin. Color partially leached from shell. Umbilicus moder-
ately widely open, U-shaped, last whorl decoiling more rapidly, con-
tained 3.78 times in the diameter. Aperture circular, slightly flat-
tened laterally above periphery, lip broken. Height of holotype 1.58
mm., diameter 2.86 mm.
Holotype. Fiji: Lau Group, Lakemba, Station 42,
Tambusue, west side near coast at 10 ft. elevation. Col-
lected by H. S. Ladd on August 8, 1934. BPBM 167222.
Range. Lakemba, Lau Group, Fiji.
Paratypes. Lakemba (2 specimens, AIM, col-
lected June 30, 1977, by W. Cernohorsky): Tambusue,
west side near coast (Station 42) at 10 ft. elevation (3
specimens, BPBM 167222); west end of Toumba
(=Tubou) (Station 97) to % mile away at 15 ft. eleva-
tion (10 specimens, BPBM 180101).
Remarks. The holotype of Sinployea inermis
lakembana is the largest and most widely umbilicated
specimen seen, but was chosen because of the clear
sculpture. Three sets were available, collected in
different years from different portions of Lakemba.
Shells taken on the west coast by Harry Ladd in 1934
(Station 42, BPBM 167222) are larger, less elevated,
and more widely umbilicated (table XXXVI) than
those found near Toumba (=Tubou) by Zimmerman
and Kondo in 1938 (Station 97, BPBM 180101). Cer-
nohorsky's 1977 specimens (Solem, 1978a) agree with
the former examples. Apparently Ladd's Station 42 is
the same as his geological Station L492, shown in Ladd
& Hoffmeister (1945, p. 99, fig. 19). It was cited as
"about 3 miles northwest of Tumbou" (Ladd &
Hoffmeister, 1945, p. 190) and thus is separated by 2.6
miles from the later collection by Kondo and Zimmer-
man. Both sets contain only adults showing marked
gerontic growth, so that the differences are not caused
by age-biased samples. Further collections are needed
to determine whether the differences are ecopheno-
typic or geographic in nature. With the umbilical vari-
ation showed by the Vanua Vatu material (table
XXXVI), I suspect that there will be no systematic sig-
nificance to this difference between the sets.
Description of soft parts. Pallial region and external body fea-
tures as in nominate subspecies. Genitalia (fig. 67f) without substan-
tive differences, more swollen spermathecal base caused by presence
of sperm packet. Shorter appearance of atrium (Y) caused by drawing
a retracted specimen with atrium and vagina folded under edge.
(Based on BPBM 180101, 2 whole and several fragmentary
examples.)
Sinployea lauensis, new species. Figure 68a-c.
Diagnosis. Shell slightly smaller than average, diameter
2.40-3.13 mm. (mean 2.76 mm.), with 3'/2-4'/4 rather loosely coiled
whorls. Apex and spire barely to moderately and evenly elevated,
last whorl descending a little more rapidly, spire protrusion about '/s
body whorl width, H/D ratio 0.493-0.581 (mean 0.530). Apical
whorls with 9-11 (mean 9.6) fine spiral cords. Postnuclear sculpture
of low, rounded, crowded, strongly protractively sinuated radial ribs,
95-119 (mean 103.9) on the body whorl, whose interstices are 1-2
times their width. Ribs/mm. 10.93-14.46 (mean 11.53). Micro-
sculpture of fine radial riblets, 2-5 between each pair of major ribs,
crossed by slightly finer and more crowded spiral riblets, with fine,
relatively crowded secondary spiral cording visible on spire and part
of the body whorl. Umbilicus narrow, V-shaped, last whorl decoiling
much more rapidly, contained 3.88-4.65 times (mean 4.24) in the
diameter, margins rounded. Whorls strongly rounded above, moder-
ately to strongly compressed laterally above periphery, aperture
subcircular, inclined about 25 from shell axis. Color reddish horn, no
darker markings.
Sinployea lauensis differs from S. inermis (fig.
66a-e) in its much less protruded spire, greater rib
frequency, and V-shaped, much more rapidly decoiling
umbilicus. Sinployea princei (fig. 65a-c) is smaller, has
fewer ribs, and the whorls are more evenly rounded
and tightly coiled, with the umbilicus decoiling more
regularly.
FIG. 68. a-c, Sinployea lauensis, new species. Station 43, Nayau, Lau, Fiji. Holotype. BPBM 167234; d-f, Sinployea navutuensis, new
species. Station 29, Navutu-i-Loma, Lau, Fiji. Holotype. BPBM 9730. Scale lines equal 1 mm. (MM).
160
SYSTEMATIC REVIEW
161
Description. Shell of average size, with slightly less than 3%
loosely coiled whorls. Apex and spire barely protruding, body whorl
descending moderately, H/D ratio 0.494. Embryonic whorls 1%,
sculpture of 9 very faint and widely spaced spiral cords. Remaining
whorls with closely spaced, narrow, rounded, strongly protractively
sinuated radial ribs, 103 on the body whorl, whose interstices are 1-2
times their width. Microsculpture of 3-6 fine radial riblets between
each pair of major ribs, crossed by lower, more crowded spiral riblets,
with some widely spaced narrow larger spiral cords. Sutures deeply
impressed, whorls shouldered above, broadly flattened between
shoulder and periphery, with slightly flattened basal margin. Um-
bilicus narrow internally, last whorl decoiling very rapidly, con-
tained 4.05 times in the diameter. Color light reddish brown. Aper-
ture subcircular, broadly flattened laterally above, inclined about
25 from the shell axis, with internal basal and columellar callus.
Columellar margin of aperture slightly reflected. Height of holotype
1.38 mm., diameter 2.80 mm.
Holotype. Fiji: Lau Group, Nayau Island, Station
43, J /2 mile inland from Nauko at 250 ft. elevation. Col-
lected at base of a limestone cliff by H. S. Ladd on
August 13, 1934. BPBM 167234.
Range. Nayau, Yangasa Levu, Namuka,
Navutu-i-Loma, and Wangava, Lau Group, Fiji.
Paratypes. Nayau: at base of limestone cliff (Sta-
tion 43) J /2 mile inland from Nauko at 250 ft. elevation
(19 specimens, BPBM 167234). Namuka: southeast
coast (Station F-3) at 50-75 ft. elevation (5 specimens,
FMNH 168089). Yangasa Levu: hillside at south end of
island (Station 30) 150 ft. inland at 40 ft. elevation (23
specimens, BPBM 167139). Navutu-i-Loma: ridge on
northeast quarter of island (Stations 28, 30) 150 yd.
inland at 150 ft. elevation (32 specimens, BPBM
167020, BPBM 167055). Wangava: northeast end of is-
land (Stations 26, 27) 100 yd. inland at 10-15 ft. eleva-
tion (7 specimens, BPBM 166955, BPBM 166968,
BPBM 166997); north end at 100 ft. elevation (38
specimens, FMNH 168092, FMNH 168101).
Remarks. Individual specimens of Sinployea
lauensis approach the measurements of S. inermis
meridionalis for single characters (table XXX), but the
different umbilical and whorl shape separate problem
specimens. Typical examples are immediately separa-
ble on a number of characters. There is considerable
similarity to the Samoan Sinployea allecta (fig. 54a-e)
in general appearance, but that species is much more
narrowly umbilicated and has less crowded radial rib-
bing.
Very little material was adult, and mean dif-
ferences between the small samples from different is-
lands are not statistically significant (table XXXI).
Sinployea navutuensis, new species. Figure 68d-f.
Diagnosis. Shell smaller than average, diameter 2.58-2.78
mm. (mean 2.66 mm.), with 4-4% normally coiled whorls. Apex and
early spire flat to barely protruding, lower spire descending slightly,
body whorl much more rapidly, spire protrusion V4 body whorl width,
H/D ratio 0.449-0.556 (mean 0.498). Apical sculpture of 9-11 (mean
10.3) narrow but prominent spiral cords. Postnuclear whorls with
narrow, crowded, fine, sharply outlined, protractively sinuated ra-
dial ribs, 121-147 (mean 136.7) on the body whorl, whose interstices
are 1-2 times their width. Ribs/mm. 16.94-17.62 (mean 17.3). Mi-
crosculpture a lattice of barely visible radial riblets, 3-6 between
each pair of major ribs, crossed by finer spiral riblets. A widely
spaced, fine sculpture of secondary spiral cording visible under an-
gled lighting only. Umbilicus broadly open, cup-shaped, regularly
decoiling, contained 3.25-3.52 times (mean 3.38) in the diameter,
margins rounded. Whorls strongly rounded, slightly compressed
laterally above periphery and on basal margin, aperture circular,
compressed laterally, inclined about 20 from shell axis.
The nearly flat apex, slight spire protrusion, wide
umbilicus, and very crowded radial ribbing separate
Sinployea navutuensis from the other Fijian species.
Sinployea lauensis (fig. 68a-c) has a much narrower
umbilicus, fewer and less crowded radial ribs, more
loosely coiled whorls, and a wider body whorl. Sin-
ployea godeffroyana (fig. 69a-c) is much smaller, much
more elevated, has a narrower umbilicus, and even
more crowded radial ribs. Sinployea kusaieana (fig.
63a-c) from the Caroline Islands is very similar in size
and shape, but differs in having only 84-103 ribs on
the body whorl and a slightly looser coiling pattern.
Description. Shell smaller than average, with 4'/s moderately
tightly coiled whorls. Apex and early spire barely elevated, body
whorl descending moderately, H/D ratio 0.487. Embryonic whorls
Ite, sculpture of 9 narrow, prominent spiral ribs. Postnuclear whorls
with low, rounded, narrow, quite crowded, protractively sinuated
radial ribs, 142 on the body whorl, whose interstices are about 1-2
times their width. Microsculpture of 3-5 very fine riblets between
each pair of major ribs, crossed by much finer, lower, spiral riblets.
Secondary sculpture of widely spaced, rounded spiral cords visible
under good lighting. Sutures deep, whorls strongly rounded above,
compressed laterally above periphery and slightly on basal margin.
Umbilicus broadly cup-shaped, regularly decoiling, contained 3.39
times in the diameter. Color leached from shell. Aperture ovate,
compressed laterally above periphery with gently rounded margin,
inclined about 15 from the shell axis. Height of holotype 1.25 mm.,
diameter 2.57 mm.
Holotype. Fiji: Lau Group, Navutu-i-Loma, Sta-
tion 29, on ridge 150 yd. inland at 100 ft. elevation.
Collected dead among limestone outcrops by H. S. Ladd
on July 24, 1934. BPBM 9730.
Range. Navutu-i-Loma and Oneata, Lau Group,
Fiji Islands.
Paratypes. Navutu-i-Loma: on ridge 150 yd. in-
land (Station 29, 1934) at 100 ft. elevation (28 speci-
mens, BPBM 167103, ex BPBM 167020, ex BPBM
167055). Oneata: 1 mile north of Dakuiloa, 1 A-1 mile
inland (Station 99, 1938) at 25-50 ft. elevation (1
specimen, BPBM 180160).
Remarks. The shell from Oneata (BPBM 180160)
is juvenile, height 0.82 mm., diameter 1.81 mm., H/D
ratio 0.455, whorls 3Vz, umbilicus 0.58 mm., D/U ratio
3.09, and may not belong to this species. It is closer to
S. navutuensis than to any other Lau species.
Only four examples were adult, the remaining 24
being juvenile or broken. The secondary spiral cording
is visible only under ideal angular lighting. It is widely
spaced and distinctly more prominent than the radial
microribbing.
Sinployea adposita (Mousson, 1870). Figures 67g,
69d-f.
Patula adposita Mousson, 1870, J. de Conchyl., 18, pp. 119-120, pi.
7, fig. 8 Oneata, Lau Group, Fiji Islands; Garrett, 1887a, Proc.
Zool. Soc. London, 1887, p. 178.
a be
FIG. 69. a-c,Sinployeagodeffroyana, new species. Station 133, Nandarivatu, 2,300-2,500 ft., Viti Levu, Fiji. Holotype. BPBM 179080; d-f,
Sinployea adposita (Mousson). Station 100, Oneata, Lau, Fiji. BPBM 180177. Scale lines equal 1 mm. (a-c, MM; d-f, SH).
162
SYSTEMATIC REVIEW
163
Helix adposita (Mousson), Pfeiffer, 1876, Monog. helic. viv., 7, p.
166.
Helix iPatula) adposita (Mousson), Tryon, 1887, Man. Conchol., (2)
3, p. 41, pi. 8, figs. 58-59.
Endodonta (Champa) adposita (Mousson), Pilsbry, 1894, Man.
Conchol., (2) 9, p. 35.
Charopa adposita (Mousson), Gude, 1913, Proc. Malacol. Soc. Lon-
don, 10 (5), p. 330; Germain, 1932, Ann. Inst. Ocean., 12 (2), p.
45.
Diagnosis. Shell large, diameter 2.63-3.75 mm. (mean 3.29
mm.), with 4-4% rather tightly coiled whorls. Apex and spire moder-
ately to strongly and evenly elevated, body whorl descending much
more rapidly, spire protrusion about l k-Yt body whorl width, H/D
ratio 0.538-0.713 (mean 0.603). Apical sculpture of 12-14 (mean
13.0) rather fine spiral cords. Postnuclear sculpture of narrow, fine,
strongly protractively sinuated radial ribs, 36-74 (mean 53.3) on the
body whorl, whose interstices are 3-6 times their width. Ribs/mm.
3.35-7.23 (mean 5.28). Microsculpture of extremely fine radial rib-
lets, usually more than 10 between each pair of major ribs, crossed by
barely visible spiral riblets. Secondary spiral cords much more prom-
inent than microradials, interstices about 3-4 times their width.
Umbilicus very narrow, U-shaped, not or only slightly decoiling, con-
tained 7.87-17.0 times (mean 11.9) in the diameter, margins
rounded. Whorls strongly and almost evenly rounded, very slightly
compressed laterally above periphery and on basal margin, aperture
ovate, inclined about 30 from shell axis.
Sinployea adposita differs from all other Fijian
species in its very narrow umbilicus and presence of
relatively regularly spaced, major radial ribbing.
Other Fijian species of similar size have either widely
open umbilici or if the umbilical range approaches that
of S. adposita, they have lost the major radial ribbing
(S. recursa, fig. 64b) or have an angulated periphery (S.
angularis, fig. 64e).
Description. Shell relatively large, with slightly less than 4%
moderately loosely coiled whorls. Apex and spire moderately and
evenly elevated, last whorl not descending much more sharply, H/D
ratio 0.622. Apical whorls l'/2, sculpture of about 12 fine spiral ribs
with a secondary sculpture of much smaller, more closely spaced
radial riblets. Remaining whorls with thin, protractively sinuated
radial ribs, 49 on the body whorl, whose interstices are about 3-5
times their width. Microsculpture a lattice of very fine radial ribs
crossed by finer, much more crowded spiral riblets with a secondary
sculpture of much more prominent, moderately closely spaced spiral
cords. Sutures moderately deep, whorls strongly rounded above, very
slightly flattened laterally above periphery. Umbilicus very narrow,
U-shaped, not decoiling, contained 13.60 times in the diameter. Color
light reddish horn, without darker flammulations. Aperture ovate,
inclined about 20 from the shell axis. Height of lectotype 1.81 mm.,
diameter 2.91 mm.
Lectotype. Fiji: Lau Group, Oneata. Collected by
Dr. Graeffe. JDCP 5246.
Range. Oneata, Munia, Lakemba, Aiwa, Mothe,
Tuvutha, Karoni, Nayau, Yangasa Levu, Lau Group,
Fiji Islands.
Paratypes. Zurich, FMNH 116979, JDCP.
Material. Oneata: southwest end of village (Sta-
tion 100) V* mile inland at 100 ft. elevation (30 speci-
mens, BPBM 180177-80). Lakemba (1 specimen,
BPBM 180117; 30 specimens. AIM, FMNH, collected
June 24-30, 1977, by Walter Cernohorsky): west end
Toumba (Station 97) 10 ft. to V 4 mile inland at 15 ft.
elevation (26 specimens, BPBM 180099-102). Munia:
Hysik's plantation (Station 64), 50 yd. inland, 100-600
ft. elevation (1 specimen, BPBM 179294); northeast
middle of island (Station 65), back of Hysik's home, 2
mile inland at 600-900 ft. elevation (5 specimens,
BPBM 179322). Nayau: % mile north of Liku Village
(Station 101) 100 yd. inland at 30-500 ft. elevation (14
specimens, BPBM 180260); Nauko (Station 43) Vz mile
inland at 250 ft. elevation (4 specimens, BPBM
167235). Yangasa Levu: south end Yangasa Levu (Sta-
tion 30) 150 ft. elevation (67 specimens, BPBM
167140). Aiwa: Vataniyambia, southwest quarter of is-
land (Station 40) 50 yd. inland at 25 ft. elevation (7
specimens, BPBM 167210). Karoni (100 specimens,
AIM, FMNH, collected July 4, 1977, by Walter Cer-
nohorsky): slope of peak at 90 ft. elevation (9 speci-
mens, BPBM 78604-5); Mothe (1 specimen, AIM, col-
lected July 7, 1977, by Walter Cernohorsky). Tuvutha:
(Station F-7) west coast at 100 ft. elevation (6 speci-
mens, FMNH 168132).
Remarks. Specimens of Sinployea adposita show
considerable variation in spacing of the major radial
ribs and some variation in shell height. Mainly dead
material in small sets was available.
Shells from Munia and the type set from Oneata
(table XXXVII) are distinctly Hatter with a lower H/D
ratio than those from the remaining islands. Possibly
there is subspeciation, but since most material was
both dead and juvenile, such separation is premature.
Shells collected from Oneata Island in 1938 are
unusual in having the radial ribbing extremely widely
spaced, with 37-57 (mean 42.7) ribs on the body whorl.
Those from Munia average 56 ribs on the body whorl;
one unworn adult from Lakemba had 68; on Yangasa
Levu they averaged 70.5; the single adult from Karoni
Islet has 72; and on Nayau shells from two populations
show the greatest extremes. Rib counts were not made
on the 1977 collections from Lakemba and Karoni
(Solem, 1978a). Specimens from % mile north of Liku
collected in 1938 have an average of 76.3 ribs on the
body whorl, whereas the one adult collected at Nauko
in 1934 has only 45 ribs, and the juveniles also have
very widely spaced ribbing. This cannot be interpreted
as meaning that the spacing of the ribbing necessarily
varies seasonally since the two localities on Nayau
yielded the quite distinct species Vatusila kondoi and
nayauana. The variation in rib spacing shows no geo-
graphic pattern, and the populations show practically
no variation in adult size (table XXXVII). The shells
from Yangasa Levu, Mothe, and Munia have slightly
narrower umbilici, but a minor size change produces a
large numerical shift in this range of measurements.
Description of soft parts. Fragmentary extracted individuals
yielded partial genitalia and one pallial region. Latter typical, kid-
ney 1.45 mm. in greatest length, rectal arm slightly longer than
pericardia!, arms of ureter widely separated by strip of lung roof.
Genitalia without unusual external features. Penis (fig. 67g, P)
about 1.65 mm. long, internally with typical vergic papilla and small
stimulatory pad.
TABLE XXXVII. - LOCAL VARIATION IN SINPLOYEA ADPOSITA AND S. IRREGULARIS.
NUMBER OF
SPECIMENS
DIAMETER
H/D RATIO
Oneata
Zurich
9 45.3*2.17
(36-54)
1.87*0.057
(1.62-2.09)
3.2310.087
(2.95-3.64)
0.57910.0080
(0.544-0
.607)
Sta. 100, 1938
BPBM 180177-80
9 42.7*3.04
(37-57)
2.03*0.042
(1.85-2.22)
3.3010.053
(3.11-3.64)
0.61510.0080
(0.587-0
.656)
Munia
Sta. 64-5, 1938
BPBM 179322, -294
4 56.32.33
(52-60)
1.80*0.136
(1.56-2.12)
3.09+0.192
(2.72-3.51)
0.58110.0105
(0.533-0
.603)
Lakemb a
Sta. 97, 1938
BPBM 180099
1
2.090.099
(1.89-2.19)
3.2810.057
(3.18-3.38)
0.63610.0329
(0.575-0
.687)
Sta. 98, 1938
BPBM 180102, -17
6 68
2.09*0.045
(1.95-2.22)
3.42*0.084
(3.05-3.64)
0.613+0.0139
(0.574-0
.650)
Yangasa Levu
Sta. 30, 1934
BPBM 167140
5 70.50.50
(70-71)
1.97*0.036
(1.85-2.05)
3.21*0.053
(3.08-3.34)
0.614+0.0098
(0.594-0
.649)
Nayau
Sta. 101, 1938
BPBM 180260
4 76.3*6.18
(67-88)
1.90*0.109
(1.62-2.12)
3.2710.092
(3.01-3.44)
0.581+0.0205
(0.538-0
.616)
irregularis
1 90 + 072
2oc+ri HAR
666*0 020
Zurich
BPBM 7982
-]
(1.71-2.20)
1. 79*0. 058
. o_)iU . U4o
(2.67-3.08)
2 78+0.041
(0.613-0
0. 643+0.014
.764)
(1.56-1.99)
(2.65-2.98)
(0.588-0
.705)
Sta. 133, BPBM
179076, -9 2300-
2500'
18
1.65*0.026
(1.51-1.91)
2.900.022
(2.70-3.09)
0.576+0.0091
(0.528-0,
.631)
Sta. 118, BPBM
178897 3000-
3600'
6
1.71*0.043
(1.61-1.91)
2.9710.030
(2.89-3.09)
0.575+0.0016
(0.544-0
.617)
Sta. 139, BPBM
179157-8 2500-
2900'
2
2.040.281
(1.76-2.32)
3.23+0.315
(2.91-3.54)
0.629+0.0262
(0.602-0
.655)
WHORLS
UMBILICUS
D/U RATIO
APICAL CORDS
adposita
Oneata
Zurich
4 l/4+(4-4 1/2)
0.27*0.012
(0. 23-0.33'"
12.210.56
(10.3-14.8)
Sta. 100
4 l/2-(4 1/4-4 3/4)
0.30*0.021
(0.20-0.36)
11.410.78
(8.82-16.7)
13
Munia
Sta. 64
4 3/8-(4-4 1/2)
0.24*0.043
(0.16-0.33)
13.811.72
(10.0-17.0)
Lakemba
Sta. 97
4 5/8-K4 1/2-4 7/8)
L 1/9 ( L I/A L *\ 1 $.\
0.30
36 + 031
11.010.17
(10.7-11.3)
13
S ta. 98
4 LI . \Q LI 44 I)/ o )
(0.30-0.46)
9 . 6410 . 66
(7.85-11.5)
Yangasa
c f. a on
L T/ft ( L 1/R L 1/71
21 + 080
15 2*0 63
b ta * JU
H Jf Q \^ LI o 4 LI )
(0.20-0.23)
(13.3-16.8)
Nayau
Sta. 101
4 3/8-(4 1/4-4 1/2)
0.31+0.095
(0.30-0.33)
10.410.19
(10.1-11.0)
irregularis
Zurich
3 7/8-(3 5/8-4)
0.58*0.019
(0.49-0.66)
4.92+0.150
(4.35-5.36)
BPBM 7982
3 5/8+(3 1/2-3 7/8)
0.58*0.021
(0.49-0.66)
4.760.159
(4.25-5.46)
18.0+1.16
(16-20)
Sta
Sta. 133
3 3/4+(3 5/8-4)
0.6910.013
(0.59-0.79)
4.2210.064
(3.79-4.77)
Sta. 118
3 3/4-(3 1/4-4)
0.680.025
(0.59-0.76)
4.3910.143
(4.13-4.88)
Sta. 139
4 1/8(3 7/8-4 3/8)
0.8110.116
(0.69-0.92)
4.0110.184
(3.82-4.19)
164
SYSTEMATIC REVIEW
165
Sinployea irregularis (Garrett, 1887). Figures 65d-f,
67a-d.
Patula irregularis Garrett, 1887, Proc. Zool. Soc. London, 1887, p.
179 Vitu Levu, Fiji Islands.
Patula monstrosa Ancey, 1889, Le Naturaliste, 1889, p. 71- Vitu
Levu, Fiji Islands. New name for irregularis Garrett, 1887, not
Endodonta irregularis Semper, 1874; Pilsbry, 1892, Man. Con-
chol., (2)8, p. 82.
Endodonta (Charopa) monstrosa (Ancey), Pilsbry, 1893, Man. Con-
chol., (2) 9, p. 35. Listed irregularis Garrett, 1887, as a synonym,
assuming it to be preoccupied by Endodonta irregularis Semper,
1874, now classified in Aaadonta.
Charopa monstrosa Ancey, Gude, 1913, Proc. Malacol. Soc. Lon-
don, 10 (5), p. 330, pi. 14, figs. 6a, b, c; Germain, 1932, Ann. Inst.
Ocean., 12 (2), p. 45.
Diagnosis. Shell of average size, diameter 2.50-3.52 mm.
(mean 2.87 mm.), with 3'A-4% rather tightly coiled whorls. Apex and
early spire flat to slightly elevated, lower spire descending moder-
ately, body whorl often descending abruptly near aperture, spire
protrusion variable, from 'Ao-% (usually Vs) body whorl width, H/D
ratio 0.528-0.764 (mean 0.614). Apical sculpture of 16-20 (mean
18.5) fine, prominent spiral cords, becoming sinuated on crossing
radial swellings. Postnuclear sculpture of highly variable, irregu-
larly spaced, low to high, protractively sinuated radial ribs, rarely
regular and persistent enough to count. Microsculpture of fine radial
riblets, finer and more crowded spiral riblets, and rather prominent,
widely spaced secondary spiral cords. Umbilicus relatively narrow,
V-shaped, last whorl regularly to more rapidly decoiling, contained
3.79-6.83 times (mean 4.54) in the diameter, margins rounded.
Whorls strongly rounded above and on outer margins, flattened to
weakly channeled laterally above periphery. Aperture circular, in-
clination varying from 10 to 30 depending on rate of body whorl
descension.
The flat spire, irregular ribbing, and often abrupt
body whorl descension effectively separate typical
examples of Sinployea irregularis from other Fijian
species. Sinployea inermis (fig. 66a-e) is separable by
its regular ribbing, lower apical cord count, and ele-
vated spire. Sinployea rudis (fig. 47d-f) from
Rarotonga, Cook Islands, is very similar in sculpture
and whorl contour, but is much larger (mean diameter
4.04 mm.), with only 8-12 apical cords, and has the
apex distinctly elevated. Ba humbugi (fig. 74a-c) has
the apex strongly elevated, few and very prominent
apical cords, and the umbilicus a lateral crack or
closed.
Description. Shell of average size, with 3% moderately tightly
coiled whorls. Apex and early spire fiat, last whorl descending
abruptly, H/D ratio 0.623. Apical whorls l'/2, sculpture of about 16
wavy, prominent spiral ribs, crossing low, broadly rounded radial
ribs that are stronger on the last portion of apical whorls. Postapical
sculpture of irregular, lamellate, protractively sinuated radial ribs,
greatly reduced on body whorl. Microsculpture of fine, irregularly
spaced, radial riblets, crossed by finer and more crowded spiral rib-
lets, plus more prominent, equally spaced spiral cords. Sutures deep,
whorls strongly rounded above, strongly flattened laterally above
periphery. Umbilicus narrow, last whorl decoiling more rapidly, con-
tained 4.25 times in the diameter. Color light yellow-brown, par-
tially leached from shell. Aperture nearly circular, somewhat de-
flected and flattened above periphery with protruded umbilical lip.
Height of lectotype 1.74 mm., diameter 2.80 mm.
Lectotype. Fiji: Viti Levu, ex Garrett, Museum
Godeffroy. BPBM 7982.
Range. Interior of Viti Levu, Fiji.
Paratypes. Zurich, FMNH 116977, BPBM 7982.
Material. Viti Levu (22 specimens, BPBM
167434, FMNH 46376, FMNH 48996, FMNH 46439):
Saganakoreva area back of Tubarua sawmill (Station
VL-2, 1965), 5 miles in from Ngaloa, Nuku District, at
950-1,000 ft. elevation (4 specimens, FMNH 153599,
FMNH 153611); Mt. Korobamba (Station 60), 2 miles
in at 1,000-1,300 ft. elevation (2 specimens, BPBM
178759); Nandarivatu (Station 129), Nukunuku Creek,
12 miles inland at 2,000 ft. elevation (1 specimen,
BPBM 179031); Nandarivatu (Station 134), 1 mile east
of Vatuthere, 12 miles inland at 2,000 ft. elevation (1
specimen, BPBM 179085); Nandarivatu (Station 133),
2 miles east of Vatuthere, 14 miles inland at 2,300-
2,500 ft. elevation (31 specimens, BPBM 179076-9);
Navai (Station 139), 1 mile west of Nandarivatu, 16
miles inland at 2,500-2,900 ft. elevation (2 specimens,
BPBM 179157-8); northwest of Nangaranambulata
Mt. (Station 122), 10 miles inland at 2,700-3,200 ft.
elevation (1 specimen, BPBM 178955); Nandarivatu
(Station 118), 10 miles inland at 3,000-3,600 ft. eleva-
tion (7 specimens, BPBM 178897-8); 500 ft. ridge (Sta-
tion F-12) near main highway, Yarawa, southeast re-
gion (12 specimens, FMNH 168309, FMNH 168311);
near Tau Village (Station F-14), 2 miles inland at 500
ft. elevation (1 specimen, FMNH 168336); Nan-
darivatu (Station F-16), ridge slope at 2,500 ft. eleva-
tion (2 specimens, FMNH 168363, FMNH 168382).
Ovalau: upper Totoga Creek (Station F-10), in from
Levuka (6 specimens, FMNH 168240, FMNH 168250).
Remarks. We do not know where on Viti Levu
the original specimens, presumably collected by Dr.
Graeffe, were obtained. The types have a high H/D
ratio produced by very sharp descension of the last
whorl or whorl and one-half. Shells nearest to these,
but distinctly less elevated and with a wider umbilicus
(table XXXVII), were obtained by members of the
Lapham Expedition at 2,500-2,900 ft. elevation near
Navai lumber camp on the slopes of Mt. Victoria (Sta-
tion 139). Shells from the vicinity of Nandarivatu (Sta-
tions 118, 129, 133, 134) and northwest of Mt. Nan-
garanambulata (Station 122) are much less elevated
(table XXXVII) and have the radial sculpture greatly
reduced in size. Shells from Saganakoreva also have
relatively low H/D ratios and wide umbilici. The im-
portance of these variations is unknown. The two
specimens from near the top of Mt. Korobamba (Sta-
tion 60) are as flat as the Nandarivatu shells, but the
radial ribs are slightly more prominent and regularly
spaced with very high periostracal extensions. Much
more material from the interior of Viti Levu is needed
in order to access the range of variation in S. ir-
regularis.
A single scalariform example, with the last third
of the body whorl completely detached, was seen. It has
not been included in the measured material.
Description of soft parts. Foot and tail retracted in all available
specimens, general appearance typical.
Body color light yellow-white, no darker markings.
Pallial region extending % whorl apically. Lung roof clear, with-
out granulations. Kidney bilobed, maximum observed length 1.8
166
SOLEM: ENDODONTOID LAND SNAILS
mm., rectal lobe distinctly longer than pericardial. Ureter typical, no
lung roof visible between arms. Heart % length of pericardial kidney
arm, lying parallel to hindgut.
Ovotestis typical, occupying '/a whorl above stomach apex, one
clump of acini lying parallel to whorl sides. Hermaphroditic duct
typical. Albumen gland (fig. 67a, GG), talon (GT), and carrefour (X)
without unusual features (fig. 67b). Prostate (DG) and uterus (UT)
typical.
Vas deferens (VD) very slender for entire length. Epiphallus (E)
with head greatly swollen, tapering rapidly to a narrow neck enter-
ing penial head lateral to insertion of penial retractor (PR), which is
very short and arises from diaphragm. Penis (P) with upper section
enormously swollen, about 0.8-1 mm. long, basal shaft slightly
shorter and very narrow. Internally (fig. 67c-d) with vergic papilla
greatly enlarged, stimulatory pad greatly reduced in prominence.
Atrium (Y) long and narrow.
Free oviduct (UV) wide, almost equal in diameter to penis.
Spermatheca (S) with very slender shaft, until just before joining free
oviduct, head elongately oval, slender. Vagina (V) tapering rapidly
to atrium.
(Based on BPBM 178897, 4 fragmentary examples.)
Sinployea godeffroyana, new species. Figure 69a-c.
Diagnosis. Shell very small, diameter 1.95-2.32 mm. (mean
2.12 mm.), with 3%-4 normally coiled whorls. Apex and spire
slightly and evenly elevated, last whorl descending slightly, spire
protrusion about l h body whorl width, H/D ratio 0.557-0.619 (mean
0.595). Apical whorls with 8-11 (mean 9.67) very fine spiral cords.
Postnuclear sculpture of very fine, narrow, low, protractively
sinuated radial ribs, 85-152 (mean 117.7) on the body whorl, whose
interstices are 1-4 times their width. Ribs/mm. 20.99-38.03 (mean
28.36). Microsculpture of very fine radial riblets, 2-5 between each
pair of major ribs, crossed by much finer and more crowded spiral
riblets, with a secondary sculpture of low and rather crowded spiral
cording. Umbilicus narrow, V-shaped, regularly decoiling, contained
4.11-6.30 times (mean 5.26) in the diameter, margins rounded.
Whorl contours typical, strongly rounded above, flattened laterally
above periphery and weakly flattened on basal margin, aperture
inclined about 10 from shell axis.
Small size and very crowded radial ribbing distin-
guish Sinployea godeffroyana from other Fijian species.
Sinployea angularis (fig. 64d-f) is similar in size but
has less than 50 radial ribs, an angulated periphery,
and markedly elevated spire.
Description. Shell very small, with 3% moderately loosely
coiled whorls. Apex and spire only slightly elevated, last whorl
barely descending, H/D ratio 0.557. Apical whorls 1%, sculpture of 8
widely spaced, relatively low spiral ribs. Remaining whorls with low,
rounded, crowded, protractively sinuated radial ribs, 152 on the body
whorl, whose interstices are 1-2 times their width. Microsculpture of
2-5 radial riblets between each pair of major ribs, crossed by very
low and inconspicuous crowded spiral ribs, with a secondary
sculpture of narrow, widely spaced spiral cords. Sutures relatively
shallow, whorls strongly rounded above, broadly flattened laterally
above periphery and on base of shell. Umbilicus narrowly V-shaped,
last whorl decoiling slightly more rapidly, contained 4.11 times in
the diameter. Color light yellow-brown without darker flammula-
tions. Aperture subquadrangular, flattened laterally above
periphery and on basal margin, inclined about 10 from shell axis.
Height of holotype 1.28 mm., diameter 2.30 mm.
Holotype. Fiji: Viti Levu, Station 133, Nan-
darivatu, 2 miles east of Vatuthere at 2,300-2,500 ft.
elevation. Collected in damp forest by Yoshio Kondo on
September 9, 1938. BPBM 179080.
Range. Viti Levu and satellite islands, Fiji.
Paratypes. Viti Levu: Mt. Korobamba (Station
63), in dense forest at 1,000 ft. elevation (1 specimen,
BPBM 178794); Nangava Island (Station 151), off
south coast of Viti Levu (1 specimen, BPBM 179256).
Remarks. The three specimens associated here
as Sinployea godeffroyana agree in general shape and
form, but diverge somewhat in sculpture. The shell
from Nangava Island (BPBM 179256) has the sculp-
ture much more widely spaced, and the umbilicus is
much narrower, with the last whorl less expanded. The
single specimen from Mt. Korobamba (BPBM 178794)
was injured several times during its growth. The nar-
rower body whorl and umbilicus may be a result of
repaired injuries rather than indicating specifically
important differences.
Until more material is available I prefer to treat
these as members of one species, although the Nan-
gava Island shell may belong to a separate taxonomic
entity.
MELANESIAN Sinployea
Without dissection of the Solomon and Bismarck
species, their classification in Sinployea rather than in
the most generalized group of Pilsbrycharopa (see
Solem, 1970a) will be somewhat arbitrary. In general
appearance these species have more similarities to
Sinployea, but either classification could be correct. I
have taken this opportunity to provide expanded diag-
noses of the Bismarck Archipelago S. novopommerana
and S. descendens, plus new descriptions of S. nissani
and S. solomonensis.
These species share monochrome coloration,
rather tight coiling, wider than usual umbilici (except
S. solomonensis), average ribbing, rather low H/D
ratios, and are the smallest in average size for any area
sampled (table XXXVIII). OnlyS. solomonensis andS.
euryomphala even approach average size for the genus.
The other four are among the seven smallest Sinployea
species. Only the Fijian S. angularis and S. godef-
froyana plus the Ellice Island S. ellicensis nukulae-
laeana are slightly smaller.
Dissection of the penial complex in S. euryomphala
confirmed my impression that Sinployea extends into
the New Hebrides. Dissection of some Andrefrancia
from New Caledonia (unpublished data) show that
they are generically distinct. The only dissected
Pilsbrycharopa from New Guinea is not one of the
generalized species, so the closeness of relationship is
uncertain.
Less detailed study has been made on this mate-
rial, but a review is included here rather than making
a separate report. Obviously collections are fragmen-
tary and do not indicate actual distributions, but only
the transitory passages of malacological collectors or
friends of malacologists. Both Bismarck species came
from river drift, although they may be sympatric. All
three known Solomon Island species are allopatric,
whereas the single New Hebridean species seems to be
widely distributed in both the New Hebrides and
Banks Group. Very probably, additional taxa will be
SYSTEMATIC REVIEW
167
TABLE XXXVIII. - RANGE OF VARIATION IN MELANESIAN SINPLOYEA.
NAME
NUMBER OF
SPECIMENS
RIBS
kuntzi
(Solem)
novopomme rana
(Rensch)
descendens
(Rensch)
solomonensis
(Clapp)
euryomphala
(Solem)
38
90
10
87
102.7(93-117)
92.7(67-105)
91.7(77-103)
89
74(72-78)
RIBS/MM.
12.85
HEIGHT
1.00
15.0(13.64-16.18) 1.17(0.99-1.37)
13.09(9.61-16.94) 1.18(0.99-1.61)
12.93(11.13-14.65) 1.18(1.05-1.28)
10.31
1.38
9.43(8.93-9.94) 1.38(1.28-1.48)
DIAMETER H/D RATIO
2.15 0.466
2.18(2.06-2.30) 0.537(0.480-0.593)
2.27(1.97-2.85) 0.519(0.467-0.567)
2.27(2.14-2.53) 0.518(0.470-0.550)
2.75
0.503
2.64(2.43-2.93) 0.523(0.494-0.535)
APICAL
CORDS
SPIRE
ELEVATION
BODY WHORL
WIDTH
SP/BWW
WHORLS UMBILICUS D/U RATIO
nissa. 3 1/2 0.67 3.20 10
kuntzi 3 3/4(3 1/2-4) 0.56(0.51-0.61) 3.91(3.68-4.26) 11.5(11-12) 0.12(0.10-0.15) 0.63(0.56-0.71) 0.191(0.176-0.209)
novop. 3 3/4-(3 1/2-4 1/8) 0.65(0.53-0.92) 3.49(3.09-3.96) 10.8(10-12) 0.09(0.07-0.13) 0.69(0.61-0.76) 0.124(0.098-0.174)
desce. 4 l/2-(4-4 1/2) 0.83(0.74-0.95) 2.73(2.43-2.94) 10.6(9-14) 0.04(0.03-0.05) 0.82(0.79-0.86) 0.049(0.038-0.063)
solom. 4+ 0.64 4.29 12
0.67(0.59-0.82) 3.97(3.56-4.67) 12.0(11-13) 0.11(0.10-0.13) 0.74(0.63-0.82) 0.149(0.120-0.170)
euryo. 3 3/4+(3 1/2-4)
recorded when more adequate collections have been
made.
(Solem, 1959). Figures
Sinployea euryomphala
67i-j, 70a-c.
Mocella euryomphala Solem, 1959, Fieldiana: Zoology, 43 (1), pp.
83-85, pi. 6, figs. 7-8 (radula and penis complex), pi. 31, figs.
1-5 (shell) Brigstock Point, Espiritu Santo, New Hebrides;
Solem, 1962, Bull. Brit. Mus. Nat. Hist., Zool., 9 (5), p. 226
Norovorovo, Maewo, and Gaua, Banks Group, New Hebrides;
Solem, 1963, Verhandl. Naturf. Ges., Basel, 74 (2), p. 163 Malo
Island off Espiritu Santo, New Hebrides.
Range. Espiritu Santo, Maewo, Gaua, Vanua
Lava, New Hebrides.
Material. Espiritu Santo: Brigstock Point (4
specimens, MCZ 186828, FMNH 54906, FMNH
109493). Maewo (1 specimen, FMNH 109384). Vanua
Lava (5 specimens, AIM, AMS).
Remarks. Use of the New Zealand genus Mocella
as a repository for this species was based on a desire to
avoid description of a new genus on inadequate data.
Dissection of Mocella (unpublished data) shows that it
is not closely related.
Very few specimens have been seen from any
single locality. I have not found any significant
differences and doubt that any separation into local
races has occurred. A description of the anatomy is
presented below, but illustrations were prepared only
for the penial complex (fig. 67i-j).
Description of soft parts. Foot bluntly rounded anteriorly,
slightly tapering and bluntly rounded posteriorly. Pedal grooves
very high on foot, uniting over tail, suprapedal slightly weaker than
pedal. No caudal foss, caudal horn, or middorsal groove. Foot sole
undivided and without corrugations. Slime network finely textured,
rectangular.
Body color pale yellow in preservative, ommatophores black.
Mantle collar short, thick, edge tapering. Pneumostome masked
by a very small anterior left mantle lobe. No other accessory lobes or
shell laps. Anus opening above and slightly anterior to external
ureteric pore, which leads into a narrow V-groove, the urinary
chamber, passing through pneumostome and mantle collar.
Pallial region typical of subfamily. Kidney bilobed, rectal lobe '/;>
longer than pericardia! and lapping onto hindgut. Ureter sigmure-
throus, not expanded at any place, with strip of lung roof between
arms. Hindgut continuing along parietal-palatal margin well apicad
of pallial cavity. Heart elongated, slightly more than M> length of
pericardial kidney lobe. Lung roof clear, without color patches.
Ovotestis a single clump of finger-like acini buried in digestive
gland well above stomach-intestine reflexion. Hermaphroditic duct
an iridescent muscular tube, greenish in tone, narrowing at base of
albumen gland, then running up to enter duct of talon at midpoint.
Head of talon a white bulb, duct slender, opening directly into
prostate-uterus head. Latter area torn in dissection, greatly distorted
because of contraction, not worked out as to detailed structure.
Vas deferens a very slender tube passing up along lower half of
penis, entering expanded tube of epiphallus above midpoint of penis.
Epiphallus (fig. 67i, E) an expanded tube with clearly defined lon-
gitudinal pilasters, reflexed, with penial retractor (PR) inserting
below point of reflexion (? artifact of contraction) and arising on
diaphragm. Penis (P) showing two bulges externally. Apical portion
with tiny verge (PV), the walls immediately below expanded to form
a glandular collar (fig. 67j), lower part with a large glandular
stimulatory papilla (PP) and a peculiar U-shaped pocket (PC) with
thin muscular walls. Base of penis narrowing to atrium (Y) with
weak pilasters present.
Free oviduct muscular, thick, with terminal glandular collar
above junction with spermatheca. Latter with grossly expanded,
thin-walled glandular base, narrowing to a very fine tube about mid-
point of free oviduct. Position of spermathecal head not observed.
Vagina very short and thin-walled.
168
SOLEM: ENDODONTOID LAND SNAILS
Digestive and free muscle system not studied. Buccal mass lost
in dissection.
The single specimen available for dissection was retracted far
within the shell and proved quite difficult to dissect. Destruction of
the shell was mandatory, although the fragments have been saved.
Diameter of the specimen was 2.27 mm. Radular elements of this
species were figured previously (Solem, 1959a, pi. 6, fig. 7).
1923). Figure
FIG. 70. a-c, Sinployea euryomphala (Solem). Holotype of
Mocella euryomphala Solem. Under logs at Brigstock Point, Espiritu
Santo, New Hebrides. UMMZ 186042. Scale line equals 1 mm.
(Solem, 1959a, pi. 31, fig. 1-3). (EJP).
Sinployea solomonensis (Clapp,
71a-e.
Endodonta (Charopa ) solomonensis Clapp, 1923, Bull. Mus. Comp.
Zool., Harvard, 65, p. 378, figs. 21-24 Ugi, Solomon Islands.
Mocella solomonensis (Clapp), Solem, 1959, Fieldiana: Zoology, 43,
p. 83: Solem, 1960, J. Malacol. Soc. Australia, 4, p. 41, pi. 4, figs.
1-5.
Description. Shell of average size, with a little more than 4
normally coiled whorls. Apex and early spire slightly emergent, body
whorl descending a little more rapidly, H/D ratio 0.503. Apical
whorls 1%, sculpture of 12 rather small spiral cords. Postnuclear
whorls with prominent, rounded, somewhat protractively sinuated
radial ribs, 89 on the body whorl, whose interstices are 2-4 times
their width. Ribs/mm. 10.31. Microsculpture mostly eroded, in places
visible as a lattice of fine radial riblets crossed by even finer and
more crowded spiral riblets. Sutures deep, whorls somewhat shoul-
dered above, flattened laterally above periphery with slightly flat-
tened basal margin. Color light reddish yellow-horn without darker
markings. Umbilicus V-shaped, regularly decoiling, contained 4.29
times in the diameter. Aperture subcircular, slightly flattened later-
ally above periphery, inclined about 15 from the shell axis. No aper-
tural barriers. Height of holotype 1.38 mm., diameter 2.75 mm.
Holotype. Solomon Islands: Ugi Island. Collected
by William Mann. MCZ 36838.
Range. Ugi, Solomon Islands.
Remarks. A proportionately thinner body whorl
and slightly more crowded radial ribbing are the most
obvious features separating this species from Sinployea
euryomphala.
Sinployea kuntzi (Solem, 1960). Figure 72a-c.
Mocella kuntzi Solem, 1960, J. Malacol. Soc. Australia, 4, pp.
41-42, pi. 5, A, figs. 1-3 Station FLW 38, peninsula east of
Halavo, Florida Island, Solomon Islands.
Range. Florida Island, Solomon Islands.
Material. Florida Island: peninsula east of
Halavo (5 specimens, FMNH 54905, MCZ 186827).
Remarks. Data on variation are presented in
Solem (1960). Measurements in Table XXXVIII are
based upon material measured recently and thus differ
slightly from the data recorded in the original descrip-
tion. Differences from S. nissani are covered in the re-
marks under that species.
Sinployea nissani (Dell, 1955).
Gyropena nissani Dell, 1955, Pacific Science, 9, p. 328, fig. 1, e-
g near Tangalan Plantation, Nissan Id., Northern Solomons.
Mocella nissani (Dell), Solem, 1960, J. Malacol. Soc. Australia, 4,
pp. 41, 43.
Description. Shell very small, with 3'/2 moderately loosely
coiled whorls. Apex barely emergent, spire flat, last whorl not de-
scending, H/D ratio 0.466. Apical whorls 1%, sculpture partially
eroded with traces of about 10 relatively widely spaced spiral cords
remaining. Postnuclear whorls with prominent, rounded, protrac-
tively sinuated radial ribs, 87 on the body whorl, whose interstices
FIG. 71. Sinployea solomonensis (Clapp). Holotype of Champa solomonensis Clapp. Ugi Island, Solomon Islands. MCZ 36838: a-c, entire
shell; d, detail of partly worn apical sculpture, greatly enlarged; e, detail of microsculpture, greatly enlarged, absence from tops of major ribs
caused by mechanical wear. Scale line equals 1 mm. (Solem, 1960, p. 42, pi. 4). (HJW).
169
170
SOLEM: ENDODONTOID LAND SNAILS
--
FIG. 72. a-c, Sinployea kuntzi (Solem). Holotype of Mocella
kuntzi Solem. From jungle one mile behind Halavo, Florida Island,
Solomon Islands. UMMZ 184473. Scale line equals 1 mm. (Solem,
1960, p. 44, pi. 5, fig. A). (EJP).
are 2-4 times their width. Ribs/mm. 12.85. Microsculpture of fine
radial riblets, 4-7 between each pair of major ribs, with much finer,
more crowded spiral riblets and a relatively prominent secondary
sculpture of widely spaced spiral cording. Sutures deep, whorls
evenly rounded above, slightly flattened laterally above periphery
with gently rounded basal margin. Color uniform yellowish brown,
slightly faded. Umbilicus widely open, broadly U-shaped, regularly
decoiling, contained 3.20 times in the diameter. Aperture ovate, lip
badly broken. Height of holotype 1.01 mm., diameter 2.16 mm.
Holotype. Solomon Islands: Nissan Island, near
Tangalan Plantation. Collected in leaf mould by R. K.
Dell in June, 1944. DMW 2513.
Range. Nissan Island, Solomon Islands.
Remarks. As indicated by Solem (1960),
Sinployea nissani is quite closely related to M. kuntzi
from Florida Island. The latter is a distinctly higher
shell (mean height 1.17 mm.), with much narrower
umbilicus (mean D/U ratio 3.91). Even though the type
of S. nissani is subadult, the umbilical width is greater
than inS. kuntzi. Although adult descension of the last
part of the body whorl would have increased the H/D
ratio of S. nissani, presumably the more rapid decoil-
ing of the last umbilical section would have further
widened the umbilical opening, increasing the
difference. They are distinct species.
Sinployea novopommerana (I. Rensch, 1937). Fig-
ure 73a-c.
Charopa novopommerana I. Rensch, 1937, Arch. f. Naturgesch.,
n.f., 6 (4), pp. 589-590, figs. 33-34 (shell and radular teeth)
Karlei, Malkong-Bach, and Nangurup, New Britain, Bismarck
Archipelago.
Diagnosis. Shell very small, diameter 1.97-2.85 mm. (mean
2.27 mm.), with 3'/fe-4Vfe normally coiled whorls. Apex and early spire
slightly elevated, rounded above, body whorl descending much more
rapidly, spire protrusion about VH body whorl width, H/D ratio
0.467-0.567 (mean 0.519). Apical sculpture of 10-12 (mean 10.8)
very fine spiral cords. Postnuclear whorls with prominent, rounded,
protractively sinuated radial ribs, 67-105 (mean 92.7) on the body
whorl, whose interstices are 2-4 times their width. Ribs/mm. 9.61-
16.94 (mean 13.09). Microsculpture of fine radial riblets, 5-10 be-
tween each pair of major ribs, slightly finer and more crowded spiral
riblets, plus rather crowded secondary spiral cording that is slightly
more prominent than microradials. Umbilicus moderately open,
V-shaped, last whorl decoiling more rapidly, contained 3.09-3.96
times (mean 3.49) in the diameter, margins rounded. Sutures very
deep, whorls strongly rounded above, flattened laterally above
rounded periphery, with evenly rounded, slightly compressed, basal
margin. Color light reddish yellow-horn without darker flammula-
tions. Aperture subcircular, flattened laterally above periphery, in-
clined about 30 from shell axis.
Sinployea novopommerana has a much less com-
pressed body whorl and narrower umbilicus than S.
descendens (fig. 73d-f). Sinployea kuntzi has a much
more elevated spire, narrower umbilicus, and less
marked lateral flattening above the periphery (fig.
72a-c).
Range. New Britain, Bismarck Archipelago.
Material. New Britain: Karlei (34 specimens,
ZMB, FMNH 146028); Malkong (4 specimens, ZMB).
Remarks. The slightly elevated apex and rather
narrow and V-shaped umbilicus immediately separate
FIG. 73. a-c,Sinployea novopommerana (Rensch). Karlei, Weite Bucht, New Britain, Bismarck Archipelago. Paratype. FMNH 146028; d-f,
Sinployea descendens (Rensch). Karlei, Weite Bucht, New Britain, Bismarck Archipelago. Paratype. FMNH 146029. Scale line equals 1 mm.
(MM).
171
172
SOLEM: ENDODONTOID LAND SNAILS
Sinployea novopommerana (fig. 73a-c) from the sym-
patric S. descendens (fig. 73d- f). The Solomon Islands.
solomonensis (Clapp, 1923) appears quite similar but is
larger, has a slightly narrower umbilicus, the whorls
are less flattened laterally above the periphery, and
the whorl count is slightly higher (fig. 71a-e).
Sinployea nissani (Dell, 1955) and S. kuntzi (Solem,
1958) are smaller, much more tightly coiled shells.
Most of the available specimens were not quite
fully adult, so that the mean diameter recorded in the
diagnosis may be too small. The range is greatly ex-
tended in regard to height, diameter, and umbilical
width by the presence of one very large individual.
Without this specimen, the maximum height would be
1.33 mm., maximum diameter 2.47 mm., and
maximum umbilical width 0.74 mm. If this specimen is
a freak, there would be a more marked size differential
between S. novopommerana and S. euryomphala.
Sinployea descendens (I. Rensch, 1937). Figure
73d-f.
Champa descendens I. Rensch, 1937, Arch. f. Naturgesch., n. f., 6
(4), p. 590, fig. 35 Karlei, Malkong, and Matong, New Britain,
Bismarck Archipelago.
Diagnosis. Shell very small, diameter 2.14-2.53 mm. (mean
2.27 mm.), with 4-4Vi rather tightly coiled whorls. Apex and early
spire flat or slightly depressed below level of penultimate and first
half of body whorl, last half of body whorl descending markedly, total
spire protrusion about l ko of body whorl width, H/D ratio 0.470-0.550
(mean 0.518). Apical sculpture of 9-14 (mean 10.6) prominent, regu-
larly spaced spiral cords, whose interstices are less than twice their
width. Postnuclear sculpture of prominent, high, rounded, rather
crowded, slightly protractively sinuated radial ribs. 77-103 (mean
91.7) on the body whorl, whose interstices are 2-4 times their width.
Ribs/mm. 11.13-14.95 (mean 12.93). Microsculpture of fine radial
riblets, 5-8 between each pair of major ribs, finer spiral riblets, and
narrow, quite crowded, secondary spiral cords. Umbilicus broadly
open, cup-shaped, regularly decoiling, contained 2.43-2.94 times
(mean 2.73) in the diameter, margins slightly shouldered. Sutures
deep, whorls shouldered above and on basal margin, evenly rounded
on columellar margin, strongly compressed laterally above rounded
periphery. Color reddish horn fading to yellow-white in drift mate-
rial, no flammulations. Aperture suboval, sharply rounded above
and at basal margin, evenly rounded laterally, inclined about 10
from shell axis.
Sinployea descendens is characterized by its flat
spire, widely open cup-shaped umbilicus, and thick
body whorl. The sympatric S. novopommerana (fig.
73a-c) has a much narrower umbilicus, thinner body
whorl, distinctly elevated spire, and averages l /z whorl
less at identical size. Sinployea irregularis (fig. 65d-f)
from Fiji is very similar in appearance, but is much
larger, has reduced ribbing, and a narrow umbilicus.
Range. New Britain, Bismarck Archipelago.
Material. New Britain: Karlei (76 specimens,
ZMB, FMNH 146029); Matong (14 specimens, ZMB).
Remarks. Both Sinployea nissani and S. kuntzi
are more narrowly umbilicated, have a protruding
spire, and average one-half whorl less. Size variation
in the two samples is summarized in Table XXXVIII.
These were all drift specimens and undoubtedly origi-
nated from several populations.
Genus Ba, new genus
Shell of less than average size with only S'/s-SM; tightly coiled
whorls. Apex and spire quite elevated, body whorl descending
rapidly. Apical sculpture of about 12 very prominent spiral cords.
Postnuclear sculpture reduced to irregularity. Umbilicus closed or
barely open. Aperture without barriers. Pallial region with mar-
kedly deflected hindgut and kidney, latter squarish with tightly
compacted ureter arms. Ovotestis with one acinal clump, hermaph-
roditic duct short. Prostate-uterine region folded and compacted.
Penis with unusually complicated penial retractor insertion. Interior
of penis with large vergic papilla and circular muscle band, pocket
stimulator greatly reduced in size.
Type species. Ba humbugi, new species.
In pattern of sculpture and general appearance, So
humbugi is extremely similar to the sympatric
Sinployea irregularis (Garrett). The major conchologi-
cal differences are the extreme spire elevation, marked
whorl reduction, and umbilical closure of Ba humbugi.
Anatomically, the compaction and deflection of the
kidney and hindgut (fig. 75a) represent a major depar-
ture from the Charopidae pattern. Similar whorl re-
duction in the Caroline Island Russatus (fig. 90a) re-
quired only compaction into a square kidney without
deflection of either kidney or hindgut from the
parietal-palatal margin. The New Guinea endemic
Paryphantopsis (Solem, 1970a, p. 250, fig. 2, a) agrees
with Ba in showing kidney and hindgut deflection, but
the pattern of the kidney is quite different in that the
ureter arms become spread progressively further
apart. These are different solutions to the same prob-
lem of visceral hump compaction. Also part of this
problem is the alteration in penial retractor muscle
insertion.
The functioning penial surface structures of Ba
humbugi show simple character displacement from the
equivalent structures in Sinployea irregularis, thus
indicating phyletic closeness. Yet Ba humbugi is the
only known member of the Pacific Island Charopinae
to experiment with whorl reduction and visceral hump
compaction. Because it represents a basic departure
from the Sinployea pattern, I have chosen to give
generic level recognition despite its evident relation-
ships to S. irregularis.
The only known species, Ba humbugi, has been
found in Central Viti Levu from Mt. Nangaranam-
bulata south to a few miles inland from Ngaloa. At the
latter locality it was found under the same log as
Sinployea irregularis. Because it occupies a portion of
Viti Levu that includes the Mba District, I have chosen
the European spelling of the district for a generic
name. This was followed by an irresistible impulse to
use the specific name humbugi.
Ba humbugi, new species. Figures 74a-c, 75a-h.
Diagnosis. Shell of slightly less than average size, diameter
2.30-3.32 mm. (mean 2.72 mm.), with S'/s-S'/i tightly coiled whorls.
Apex and spire strongly and evenly elevated, body whorl descending
more rapidly, spire protrusion M)-M> body whorl width, H/D ratio
0.752-0.842 (mean 0.801). Apical sculpture of 11-13 (mean 12.0)
high and prominent spiral cords. Postnuclear sculpture of irregular
growth wrinkles and an occasional identifiable major rib with inter-
SYSTEMATIC REVIEW
173
FIG. 74. a-c,Ba humbugi, new species. Mt. Nangaranambulata,
2,700-3,200 ft, Viti Levu, Fiji. Holotype. BPBM 178954. Scale line
equals 1 mm. (MM).
gradation between these extremes. Microsculpture obscured by sur-
face irregularities, visible in some areas as consisting of fine radial
riblets, finer and more crowded spiral riblets, plus low and broadly
rounded, much larger secondary spiral cords. Umbilicus either im-
perforate (25%) or with a small lateral crack (75%), possibly latter
developing in adult stage only, since smallest example had closed
umbilicus. Sutures deep except near end of body whorl, aperture and
body whorl flattened laterally above periphery, inclined about 25
from shell axis.
Ba humbugi is immediately recognizable by its
very high spire, low whorl count, reduced radial rib-
bing, and closed or laterally cracked umbilicus. Other
Polynesian species with closed or nearly closed um-
bilici include the Samoan S. clausa and S. clista (fig.
51c, f) and the Lau Archipelago S. adposita (fig. 69f).
These species all have prominent radial ribbing and
higher whorl counts. The Lord Howe Island Mystivagor
mastersi (Brazier, 1872) is similar in shape and whorl
count but has vastly different sculpture and anatomy.
Description. Shell rather small, with 3'/2 relatively tightly
coiled whorls. Apex and spire markedly and evenly elevated, body
whorl descending more rapidly, H/D ratio 0.810. Apical whorls I'/z,
sculpture of 11 very prominent spiral cords. Postnuclear whorls with
very irregular growth wrinkles and occasional major ribs with high
periostracal extensions. Microsculpture a lattice of fine radial and
finer spiral riblets, irregular in spacing, crossed by more prominent
spiral cords. Sutures deep, whorls flatly rounded above, evenly
rounded below periphery. Umbilicus closed by expansion of basal lip
and tight coiling, with only a slight lateral chink visible. Color light
reddish yellow with dark brown, almost black, periostracal exten-
sions. Aperture ovate, flattened laterally above periphery, inclined
about 25 from shell axis. Height of holotype 2.24 mm., diameter 2.76
mm.
Holotype. Fiji: Viti Levu, Station 122, Mt. Nan-
garanambulata at 2,700-3,200 ft. elevation. Collected
in dense forest by Yoshio Kondo on September 5, 1938.
BPBM 178954.
Range. Interior of Viti Levu at 950-3,200 ft. ele-
vation, Fiji.
Paratypes. Viti Levu: top of Mt. Korobamba (Sta-
tion 60) at 1,000-1,300 ft. elevation (2 specimens,
BPBM 178758); Sanganakoreva area (Station VL-2,
1965), back of Tubarua saw mill, 950-1,000 ft. eleva-
tion, 5 miles inland from Ngaloa, Nuku District (1
specimen, FMNH 153605).
Remarks. The closed umbilicus, irregular rib-
bing, and very high spire of Ba humbugi immediately
separate it from any Pacific Island endodontid. In
many respects, the shell recalls that of the New
Caledonian Rhytidopsis minutula (Crosse, 1870) (see
Solem, 1961, p. 466). Until the latter can be dissected,
the degree of relationship will remain uncertain. The
Lord Howe Island species Mystivagor mastersi
(Brazier, 1872) has gone further toward a succineiform
shell. It differs in size (6 mm. diameter), strong color
pattern, smooth shell surface, and apical sculpture of
widely spaced radial ribs.
Spire angle varied greatly among the four speci-
mens, with the one from Saganakoreva (FMNH
153605) having a much wider angle and lower H/D
ratio (0.752). It was found under the same log as an
example of Sinployea irregularis, so that the two
174
SOLEM: ENDODONTOID LAND SNAILS
species are unquestionably sympatric. Lack of any
black markings on the body probably indicates this is a
strictly terrestrial species.
Dissection of two individuals showed major
changes in the pallial systems that resulted in generic
recognition. Changes in the genitalia involved folding
of the prostate-uterus, compaction and broadening of
the albumen gland, shortening of the hermaphroditic
duct, altering the simple clump insertion of the penial
retractor into a complex (fig. 75d-e), enfolding of the
penis-epiphallus junction, and massive reduction of the
penial pocket stimulator (fig. 75h). Except for the lat-
ter change, these are adjustments to the whorl reduc-
tion and visceral hump compaction.
Comparison of penial structures in S. irregularis
(fig. 67c-d) with those of Ba humbugi (fig. 75h) shows
enlargement of the pocket stimulator in the first and
reduction in the second.
A single sperm packet (fig. 75f-g) was taken from
FNMH 153605, which was collected December 9, 1965,
the morning after a shower ending a long drought.
Description of soft parts. Foot and tail equal in length to shell
diameter. Sole undivided, bluntly rounded and slightly tapering
posteriorly, truncated anteriorly. Pedal grooves very conspicuous,
high on foot, uniting above tail, no caudal horn or middorsal groove
developed. Slime network of large, rectangular sections on tail and
sides of foot. Head retracted in all individuals examined. Gonopore
position normal.
Body color yellow-white, no darker markings.
Mantle collar (MC) with thickened edge, no glandular extension
onto pallial roof. Anus (A) opening next to external ureteric pore
(KX), just inside pneumostome.
Pallia! region (fig. 75a) extending V 2 whorl apically. Lung roof
clear, without granulations. Kidney (K) about 1.58 mm. long,
bilobed, deflected downward from parietal-palatal margin after 0.33
mm., lobes subequal, pericardial slightly longer than rectal, trun-
cated anteriorly by ureter. Ureter (KD) compressed between kidney
lobes, apical part curved downward around tip of kidney, no lung
surface visible between kidney lobes. Heart (H) % length of kidney,
lying parallel to deflected plane of hindgut and parallel to lower edge
of mantle collar. Principal pulmonary vein (HV) very short and in-
conspicuous, curving around edge of kidney. Hindgut (HG) deflected
from parietal-palatal margin shortly after kidney apex, intestinal
loop (dotted line in fig. 75a) partly under heart.
Ovotestis (fig. 75c, G) occupying V-i whorl above stomach apex, a
single, tightly compacted clump of palmately clavate acini with short
collecting tubule. Hermaphroditic duct (GD) kinked initially, nar-
row, gradually tapering to past middle, grossly expanded, then
sharply constricting before reflexing up to talon. Albumen gland
(GG) with highly irregular surface, proportionately large, acini rela-
tively small. Talon (GT) with globular head, short neck to carrefour,
both buried in albumen gland. Prostate (DG) with a few long acini
inserting onto side of uterus, which is bifolded. Uterus (UT) bipar-
tite, folded upon itself, lower chamber enlarged and with thick, glan-
dular walls.
Vas deferens (VD) with thick, glandular walls to penioviducal
angle, narrowed abruptly and a thin-walled tube to epiphallus,
loosely bound to penioviducal angle. Epiphallus (E) % length of
penis, head bulbous, internally with Y valve and longitudinal pilas-
ters. Folded against head of penis at point of insertion. Penial retrac-
tor (PR) arising from diaphragm, about Mi length of penis, inserting
in a broad fan around epiphallus and head of penis (fig. 75d-e), with
separate strands to penis apex and inner wall of epiphallus. Penis (P)
club-shaped, about 1.8 mm. long, tapering gradually to atrium, in-
ternally (fig. 75h) with a bulbous vergic papilla, greatly enlarged
circular muscle band, and tiny stimulatory pocket pilaster. Lower
portion with weak glandular ridges into atrium. Atrium (Y) short,
without unusual features.
Free oviduct (UV) only slightly longer than vagina, equal in
diameter to penis at midsection, internally with typical circular
pilaster at spermathecal junction. Spermatheca (S) with expanded
head next to albumen gland, slender shaft next to prostate, base
section abruptly expanding at base of uterus, internally with typical
glandular pilaster and a larger sperm packet (fig. 75f-g, SP). Vagina
(V) thin-walled, same diameter as free oviduct, internally with weak
glandular pilasters.
Free muscle system typical, somewhat shortened. Right om-
matophoral retractor passing through penioviducal angle, right
rhinophoral retractor uniting before tail fan junction.
Stomach occupying % of a whorl, starting ] Ae of a whorl above
pallial cavity apex. Intestinal looping compacted into narrow zone
and protruding partly into pallial cavity area. Hindgut deflected
from parietal-palatal margin for half of length, reaching margin
near anterior end of kidney.
Digestive glands extending past ovotestis to apex, mainly along
stomach, only one strand near albumen gland. Salivary glands short,
not uniting above esophagus.
(Based on FMNH 153605 and BPBM 178954, 2 adult specimens.)
Genus Maafu, new genus
Shell relatively large, with 4-4V> normally coiled whorls. Apex
and spire flatly coiled, depressed below level of postnuclear sub-
sutural keel. Periphery strongly protruded into a cordlike keel with
prominent supra- and subperipheral sulci. Umbilicus broadly open,
cup-shaped, regularly decoiling, contained about 3 times in the
diameter. Apical sculpture with typical spiral cords. Postnuclear
sculpture with prominent, widely spaced radial ribs that are en-
larged on crossing periphery, regular microsculpture reduced, secon-
dary spiral cording narrow, rather prominent, and quite crowded. No
apertural barriers. Anatomy unknown.
Type species. Maafu thaumasius, new species.
Although the form of Maafu thaumasius is par-
tially duplicated by such Endodontidae as Austral-
donta magnasulcata (Solem, 1976b, p. 295, fig.
127a-b), several Nesodiscus, Endodonta, and many
juvenile Gambiodonta, no Pacific Island Charopidae
have a protruded keel. Only Sinployea angularis (fig.
64e) from Namuka, Lau Group, andHimeroconcha (fig.
105b, e) from the Marianas have an angulated
periphery.
Most of the greatly altered appearance seen in
Maafu thaumasius relates to the two keels. The sub-
sutural keel changes what would be a flat-spired shell
into one that has a depressed apex and early spire. The
extreme protrusion of the peripheral keel increases the
diameter and decreases the H/D ratio. Hence, the com-
paratively large size (mean diameter 3.49 mm.) and
very low H/D ratio (mean 0.365) are secondary modifi-
cations resulting from the keels. Efforts to assess the
relationships of M. thaumasius must discount the im-
portance of these size and shape shifts. The micro-
sculpture of Sinployea inermis meridionalis and S. i.
lakembana is visually very close to that of M.
thaumasius. Only the greater crowding and emphasis
of the secondary spiral cording distinguish the latter in
a gross sense. The pattern of coiling, umbilical shape,
and major ribbing is very different, but derivation of
Maafu from the generalized Sinployea inermis-S. prin-
cei stock would present no insuperable difficulties.
Fig. 75. Anatomy of the Fijian Ba humbugi, new species. Station VL-2, Tubarua, Viti Levu, Fiji. FMNH 153605: a, pallial region; b,
genitalia; c, ovotestis and hermaphroditic duct; d-e, insertion details of penial retractor (PR) on epiphallus-penis junction; f-g, spermatophore;
h, interior of apical penis region. Scale lines equal 1 mm. (SH).
175
176
SOLEM: ENDODONTOID LAND SNAILS
Besides the keels, the alteration in ribbing pat-
tern, very broad and regularly decoiling umbilicus, and
flat spire represent departures from the Sinployea pat-
tern.
Thus, generic recognition for Maafu is based on
several shifts that combine to produce a drastically al-
tered appearance. Local differentiation from a
Sinployea-type stock is almost certain, but the
structural gap is real and large.
I have chosen to honor the great Tongan leader
Ma'afu, who unified the Lau Group in the 1850- 1860s,
by naming this genus after him. The specific name
thaumasius refers to the most unusual and wonderful
shape of the shell.
Maafu thaumasius, new species. Figure 76a-c.
Diagnosis. Shell relatively large, diameter 3.19-3.98 mm.
(mean 3.49 mm.), with 3'/2-4'/8 normally coiled whorls. Apex and
spire depressed below level of subsutural keel, body whorl not de-
scending or rarely very slightly deflected, H/D ratio 0.343-0.381
(mean 0.365). Apical sculpture of 10-15 (mean 12.2) narrow but
prominent spiral cords. Postnuclear sculpture of high, broadly
rounded, rather widely spaced radial ribs, 36-55 (mean 44.1) on the
body whorl, whose interstices are about 2-5 times their width, and
which become higher and more sharply defined on crossing keels.
Ribs/mm. 3.59-4.74 (mean 4.08). Microsculpture of extremely fine
radial riblets, barely visible at 96 x magnification and too obscured
by the secondary spiral cording to be counted accurately, equally fine
spiral riblets, and much more prominent secondary spiral cording,
whose interstices are 1-3 times their width. Umbilicus broadly open,
cup-shaped, regularly decoiling, contained 2.82-3.15 times (mean
2.99) in the diameter, columellar-basal margin obtusely shouldered.
Sutures deep, whorls strongly rounded above, after nucleus with a
gradually increasing subsutural keel followed by a deep supra-
peripheral sulcus, periphery a sharply protruded cordlike keel, a
broader and shallower subperipheral sulcus, basal margin rounded
to an obtusely shouldered columellar-basal angle, walls of umbilicus
evenly rounded.
Although the form of Maafu thaumasius recalls
that of the Navutu-i-Loma endodontid Zyzzyxdonta
alata Solem (1976b, p. 466, fig. 198a-c), the larger size,
beaded and numerous apertural barriers, "winglike"
rib extensions, flat spire, and radial apical sculpture of
the latter at once separate the two species. No Polyne-
sian Charopidae have an even slightly similar shape,
and M. thaumasius cannot be confused with any other
species.
Description. Shell relatively large, with slightly less than 4%
moderately tightly coiled whorls. Apex and early spire depressed
below plane of subsutural keel on succeeding whorls, last whorl not
descending, H/D ratio 0.378. Sutures shallow, slightly denticulated
by radial ribbing. Postapical whorls with a high subsutural ridge,
becoming slightly less prominent near end of body whorl. Periphery
of body whorl greatly protruded into a ridgelike keel, denticulated by
crossing of major ribs, with a very prominent sulcus above and a less
prominent one below periphery. Embryonic whorls 1%, sculpture of
about 12 very fine, closely spaced spiral ribs (partially eroded). Post-
nuclear whorls with widely spaced, low, rounded, protractively
sinuated radial ribs, 50 on the body whorl, that denticulate the
periphery and whose interstices are 3-6 times their width. Micro-
sculpture on early whorls of very fine radial riblets, crossed by
slightly stronger secondary spiral cords. Microspiral riblets occa-
sionally visible. On later whorls the radials become obsolete and the
spirals slightly more prominent. Umbilicus broadly open, cup-
FIG. 76. a-c, Maafu thaumasius, new species. Station 43, Nayau,
Lau, Fiji. Holotype. BPBM 167233. (MM).
SYSTEMATIC REVIEW
177
shaped, regularly decoiling, contained 2.98 times in the diameter.
Color mainly leached from shell. Aperture subquadrangular with
concave outer margins and rostrate periphery, inclined about 20
from the shell axis. Height of holotype 1.48 mm., diameter 3.92 mm.
Holotype. Fiji: Lau Group, Nayau, Station 43,
hillside '/2 mile inland of Nauko at 250 ft. elevation.
Collected at base of limestone cliff by H. S. Ladd on
August 13, 1934. BPBM 167233.
Range. Nayau, Lau Group, Fiji.
Paratypes. Nayau, hillside 2 mile inland of
Nauko (Station 43) at 250 ft. elevation (64 specimens,
BPBM 167233).
Remarks. The very striking subsutural ridge
and peripheral keel immediately separate Maafu
thaumasius from all other species of Polynesian
Charopidae. It is most apt to be confused with Zyzzyx-
donta alata, a species of Endodontidae from Navutu-i-
Loma that differs in having radial apical sculpture and
apertural barriers of strong lamellae that are regularly
beaded above. In addition the major ribs of Z. alata are
prolonged into cup-shaped winglike structures when
crossing the periphery (Solem, 1976b, p. 466, fig.
198a-c). The ribs in M. thaumasius only slightly den-
ticulate the keel (fig. 76b).
Only 20% of the specimens were adult. Much of the
material was fresh, but no specimens were collected
alive.
Genus Lauopa, new genus
Shell extremely large, with about 4% normally coiled whorls.
Apex and spire flat or slightly depressed, body whorl descending
rapidly. Apical sculpture of about 22 very prominent spiral cords.
Postnuclear whorls with high, widely spaced, strongly protractively
sinuated radial ribs. Microsculpture of fine radials and finer spirals,
no secondary cording. Umbilicus widely open, cup-shaped, regularly
decoiling. Whorls flattened laterally above and slightly compressed
laterally below rounded periphery. Parietal wall with median, high
barrier, extending about '/i whorl, with abrupt anterior descension,
not expanded above. No palatal or columellar barriers. Anatomy
unknown.
Type species. Lauopa mbalavuana, new species.
Placement of this species in a distinct genus with-
out any knowledge of its anatomy is based upon elimi-
nation of alternative classifications. Key features are
the very large and numerous apical cords, presence of a
single parietal barrier, absence of secondary spiral
cording, very large size, and widely spaced radial rib-
bing. Species of Graeffedon from Samoa and Tonga are
much smaller, have many more and shorter barriers,
more crowded radial ribbing, and much narrower um-
bilici. They agree in lack of secondary spiral cording
and the flat or nearly flat apex and spire. Tuimalila
from Tonga also lacks secondary spiral cording, but
differs in its narrow umbilicus, distinctly elevated
spire, prominent and much more crowded radial
sculpture, greatly reduced apical sculpture, total ab-
sence of apertural barriers, and smaller size. Sinployea
complementaria (Mousson) from Upolu, Western
Samoa, is only 4.92 mm. in mean diameter and lacks
any apertural barriers (fig. 56b).
Possibly Lauopa may be related to the Microne-
sian genus Semperdon, but until live material can be
obtained and dissected, no affinities can be determined.
Generic recognition is given because there are major
differences from any other genus known in this region.
The name Lauopa refers to its occurrence in the Lau
Archipelago. For convenience it is associated with the
generalized Charopinae.
Lauopa mbalavuana, new species. Figure 77a-c.
Diagnosis. Shell extremely large, diameter 6.67-8.37 mm.
(mean 7.52 mm.), with 4>/4-4% normally coiled whorls. Apex and
spire flat, lower spire descending slightly, body whorl much more
rapidly, spire protrusion less than 'Ao body whorl width, H/D ratio
0.422-0.437 (mean 0.430). Apical sculpture of about 22 prominent
spiral cords whose interstices are about equal to their width. Post-
nuclear sculpture of high, prominent, protractively sinuated radial
ribs, 39 on the body whorl in subadult, whose interstices are about
5-8 times their width, becoming extremely crowded and irregular on
last third of body whorl in adult (fig. 77a). Ribs/mm, in juvenile 1.86.
Microsculpture of extremely fine and crowded spiral and radial rib-
lets, more than 20 radials between each pair of major ribs, spirals
distinctly finer. No secondary spiral cording. Umbilicus broadly
open, cup-shaped, regularly decoiling, contained 2.72-2.76 times
(mean 2.74) in the diameter, margins rounded. Whorls strongly
rounded above and on periphery, slightly compressed laterally on
upper and lower palatal margins. Parietal wall with single median,
high barrier, extending posteriorly less than % whorl, slightly ex-
panded above, with abrupt descension at anterior end. No columellar
or palatal barriers.
Tuimalila pilsbryi and T. infundibulus from
Tonga (fig. 78a-f) both approach the size of Lauopa
mbalavuana but lack the parietal barrier, have an ele-
vated spire, much finer apical cording, and more
crowded radial ribbing. All other Fijian and Polyne-
sian species are much smaller in size.
Description. Shell extremely large, with 4% moderately tightly
coiled whorls. Apex and early spire slightly depressed below penul-
timate, last whorl descending rapidly, H/D ratio 0.437. Embryonic
whorls l'/2, sculpture eroded. Remaining whorls with remnants of
protractively sinuate, low, rounded, widely spaced radial ribs whose
interstices are 3-7 times their width, becoming extremely crowded
on last third of body whorl. Microsculpture of very fine, close-set
riblets crossed by even more crowded, low spiral ribs. Sutures deeply
impressed, whorls strongly rounded above, flattened laterally above
periphery. Color leached from shell with few vague flammulations
remaining. Aperture circular, flattened laterally above periphery,
inclined 25 from the shell axis. Parietal wall with one high,
ridgelike barrier extending a little more than 3 Ae of a whorl. Um-
bilicus broadly open, cup-shaped, regularly decoiling, contained 2.72
times in the diameter. Height of holotype 3.66 mm., diameter 8.37
mm.
Holotype. Fiji: Lau Group, Vanua Mbalavu, Sta-
tion 78, a limestone hill between Valika and Mosomo
Bay, % mile inland at 200-250 ft. elevation. Collected
under stones and logs in a dense forest by Yoshio
Kondo and Elwood Zimmerman on August 9, 1938.
BPBM 179554.
Paratype. Lau Group: Vanua Mbalavu, a lime-
stone hill between Valika and Mosomo Bay, % mile
inland (Station 78) at 200-250 ft. elevation, under
stones and logs (1 specimen, BPBM 179554).
Remarks. The adult holotype of Lauopa mbala-
vuana is a rather badly worn specimen, whereas the
178
SOLEM: ENDODONTOID LAND SNAILS
FIG. 77. a-c, Lauopa mbalavuana, new species. Station 78,
Vanua Mbalavu. Lati. Fiji. Holotype. BPBM 179554. Scale line
equals 1 mm. (MMi.
juvenile subadult retains most of the apical and micro-
sculpture. It is uncertain whether this is a living or
extinct species because the better-preserved paratype
apparently was leached out from coralline rock, frag-
ments of which are cemented to the shell.
The only species of similar size, shape, and apical
sculpture are the Tongan Tuimalila infundibulus and
T. pilsbryi, neither of which have any apertural bar-
riers.
Genus Tuimalila, new genus
Shell very large, with normally to loosely coiling whorls. Apex
and spire moderately elevated. Apex with fine and crowded spiral
cords, at least in one species with a secondary intrusion of radial
swellings that appear under optical examination as primary radial
ribs. Postnuclear sculpture of prominent, large, major radials and
typical microsculpture. Umbilicus narrow, V-shaped, regularly de-
coiling. Aperture relatively large. Pallial region as in Sinp/oyea, a
narrow strip of lung roof visible between arms of ureter, kidney lobes
almost equal in size. Genitalia agreeing with Sinployea except for
nearly apical pore in the vergic papilla and presence of a well-
developed retractor muscle attached to the free oviduct head.
Type species. Tuimalila pilsbryi, new species.
Although the Upolu Sinployea complementaria
(Mousson) approaches the size of Tuimalila and several
Rarotongan species had individuals within its size
range, the two species grouped here average signifi-
cantly larger, show a different pattern of whorl incre-
ment, and have distinctive patterns of apertural open-
ing. Their apical sculpture has the spiral cords reduced
in prominence and increased in number over that
found in Sinployea, whereas at least in T. pilsbryi there
is developed a secondary intrusion of radial swellings
(fig. 2c, p. 11). These appear as radial ribs under optical
microscopic inspection at 96 x and visually mimic the
characteristic Endodontidae apical sculpture. Whether
this same feature occurs in T. infundibulus cannot be
determined until specimens with unworn apical
sculpture are obtained for study.
Anatomical features show no characters incompat-
ible with derivation from Sinp/oyea, whereas the con-
chological changes are matters of degree rather than
major shifts. Nevertheless, the change in apical
sculpture and increase in size are major departures
from the typical Sinployea pattern. The average mean
diameter of the two species in Tuimalila, 6.16 mm.,
represents a 547( jump over the median mean diameter
of Sinployea species and a 25% jump over the mean
diameter, 4.92 mm., of S. complementaria (Mousson),
the largest species in that genus. This size increase
occurred with a slight actual decrease in whorl count
and a marked increase in relative whorl widths. In
Sinployea the median mean whorl count is 4V-,
whereas Tuimalila infundibulus has 3%+ and T.
pilsbryi 4+ whorls. The result of this size increase
combined with no increase in whorl count is to alter the
growth pattern. The much more rapidly increasing
whorl cross-section combines with the higher pallial
surface to broaden the kidney (fig. 79a) and to fore-
shorten the pallial organization.
SYSTEMATIC REVIEW
179
Just as Ba represents a new experiment in altered
growth pattern by increased spire height, so Tuimallla
is a new pattern based on great increase in whorl
width. As such, both merit generic separation from
Sinployea.
The name Tuimalila commemorates the deceased
famous tortoise, Tu'i Malila, which was resident in
Tonga for many years and supposedly dated from the
visit of Captain Cook (see Robb & Turbott, 1971).
Tuimalila pilsbryi, new species, Figures 78a-c,
79a-e.
Champa radicalis Boettger, 1916 (not Mousson, 1871), Abhl.
Senckenb. Naturf. Gesell., 36 (3), p. 290 Eua, Tonga.
Diagnosis. Shell extremely large, diameter 5.88-6.99 mm.
(mean 6.64 mm.), with 3%-4% almost normally coiled whorls. Apex
and spire moderately and evenly elevated, body whorl descending
slightly more rapidly, spire protrusion about Vs-'/t body whorl width,
H/D ratio 0.495-0.606 (mean 0.556). Apical sculpture of numerous
extremely fine and regular spiral cords (fig. 2c, p. 11), with a gradual
intrusion of low, rounded radial swellings that appear as fine, major
radial ribs on last portion. Postnuclear sculpture of relatively nar-
row, prominent, V-shaped, strongly protractively sinuated radial
ribs, 74-103 (mean 91.2) on the body whorl, whose interstices are
2-4 times their width. Ribs/mm. 3.55-4.97 (mean 4.37). Micro-
sculpture of very fine radial riblets, 5-12 between each pair of major
ribs, crossed by distinctly finer and more crowded spiral riblets (fig.
2c-d). Umbilicus narrow, V-shaped, regularly decoiling, contained
4.48-6.50 times (mean 5.62) in the diameter, margins rounded.
Whorl contours and aperture typical.
Tuimalila pilsbryi from Eua (fig. 78a-c) is a more
depressed, larger shell with less descension of the body
whorl, and has lower, less sharply defined radial ribs
than does T. infundibulus (fig. 78d-f) from Vavau.
Description. Shell extremely large, with 4 somewhat loosely
coiled whorls. Apex and early spire moderately and evenly elevated,
body whorl not descending more rapidly, H/D ratio 0.529. Embryonic
whorls I :! /K, first whorl with sculpture of about 33 very fine spiral
riblets, last % whorl showing a gradual emergence of low, broadly
rounded, radial swellings, foreshadowing spire sculpture. Radial ribs
on early spire low, broadly rounded, becoming more prominent and
higher below. Radial ribs on body whorl protractively sinuated,
prominent, 94 in number, whose interstices are 2-3 times their
width. Microsculpture of very fine, closely set radial riblets with
equally spaced, smaller spiral ribs. Sutures moderately impressed,
whorls evenly rounded above, body whorl somewhat flattened later-
ally above periphery and slightly flattened on basal margin. Um-
bilicus open, V-shaped, last whorl decoiling slightly more rapidly,
contained 4.86 times in the diameter. Color light yellow-white with
numerous sinuated irregular reddish brown flammulations. Aper-
ture ovate with somewhat flattened basal margin, inclined about 20'
from shell axis. Height of holotype 3.53 mm., diameter 6.67 mm.
Holotype. Tonga: Eua, Johanssen Plantation, 2
miles inland at 385 ft. elevation. Collected by H. S.
Ladd on May 14, 1928. BPBM 87750.
Range. Eua Island, Tonga.
Paratypes. Eua (1 specimen, SMF 165341 col-
lected by E. Wolf on June 5, 1909): Johanssen Planta-
tion, 2 miles inland at 385-500 ft. elevation (5 speci-
mens, BPBM 87750, BPBM 87719, BPBM 108565);
main range on east side (Station T-22) of island at
1,000 ft. elevation in heavy forest (35 specimens,
FMNH 152378).
Remarks. Although there are a few striking
quantitative differences (table XXXIX) between
Tuimalila pilsbryi and T. infundibulus, the ribbing
character and different aperture appearances im-
mediately separate individuals of the two species. Al-
though the intrusion of secondary radial ribbing into
the apex was not seen in any specimens of T. infun-
dibulus, all specimens of the latter were so worn that
this may be present on that species also.
Living examples were common in fallen tree fern
fronds.
Great pleasure is taken in dedicating this species
to the late Henry A. Pilsbry, dean of terrestrial mala-
cologists and mentor.
Description of soft parts. Foot and tail equal to shell diameter
in length, slightly tapering posteriorly, truncated anteriorly. Sole
undivided longitudinally, transversely corrugated in preservative.
Pedal grooves typical, suprapedal distinctly weaker than pedal, both
uniting above tail, no caudal horn or middorsal groove developed.
Slime network weak, without peculiarities. Head projecting in front
of foot, ommatophores typical, eyespots large. Gonopore a broad slit,
opening behind right rhinophore and below right ommatophore.
Body color yellow-white, a faint trace of grayish color on neck,
ommatophores distinctly grayish.
Mantle collar (MO thick, without obvious lobing, a broad black
zone of color granules along inner margin, no glandular extension
into pallial roof. Anus opening just inside pneumostome, slightly in
front of external ureteric pore (KX).
Pallial region (fig. 79a) extending slightly more than % whorl
apically. Lung roof clear, no granulations. Kidney (K) about 3.0 mm.
long, bilobed, with rectal arm slightly longer, reaching across
hindgut onto parietal wall, very high at end of cavity, dished on
posterior margin. Distance from anterior end to anus 2.1 mm. Ureter
(KD) typical, a narrow strip of lung roof visible between arms that
open at a slight angle. Heart (H) more than M; length of kidney,
slightly angled to hindgut. Principal pulmonary vein (HV) un-
branched, relatively conspicuous. Hindgut (HO) without unusual
features.
Ovotestis a single clump of palmately clavate alveoli, lying
parallel to whorl sides and occupying almost '/> whorl above stomach
apex. Hermaphroditic duct iridescent, slightly kinked in middle
areas, narrowing abruptly at point where it starts along albumen
gland, reflexinp up to enter junction of talon and carrefour (fig. 79e).
Albumen gland with rather large acini, surface deeply indented by
other organs, general shape squarish. Talon (fig. 79e, GT) globular,
without a distinct shaft, joining hermaphroditic duct directly at head
of carrefour (Xl. Latter -<i diameter of talon, tapering into head of
prostate-uterus. Prostate (DG) with many very prominent and long
alveoli complexly folded against wall of uterus, opening into groove
on inner wall of uterine chambers. Prostate-uterus folded back on
itself for part of length. Uterus (UT) typically bipartite, lower
chamber with thick, glandular walls.
Vas deferens (VD) wide in diameter with rather thick, pilas-
tered, glandular walls until just before penioviducal angle, becoming
very slender, reflexing upward. Epiphallus (E) about % length of
penis, inserting into penis to one side of penial retractor, receiving
vas deferens laterally on very swollen head, lower % more slender.
Internally with Y valve and plug apically, lower portion with lon-
gitudinal pilasters running into epiphallic pore (fig. 79c). Penial re-
tractor (PR) very short, arising on diaphragm, inserting directly onto
head of penis. Penis (P) about 2.4-2.5 mm. long, club-shaped, swollen
medially, tapering slightly to atrial junction. Internally with typical
pocket stimulator producing medial bulge, apically (fig. 79c-d) with
vergic papilla and circular muscle band on wall below it. Atrium (Y)
about 'it length of penis, tapering gradually.
FIG. 78. a-c, Tuimalila pilsbryi, new species. Johanssen Plantation, Eua, Tonga. Holotype. BPBM 87750; d-f, Tuimalila infundibulus
(Hombron & Jacquinot). Lectotype ofPatula radicalis Mousson. Zoologisches Institut der Universitat Zurich. Scale lines equal 1 mm. (a-c, SH;
d-f, SG).
180
SYSTEMATIC REVIEW
TABLE XXXIX. - RANGE OF VARIATION IN FIJIAN AND TONGAN ENDEMIC GENERA.
181
NUMBER OF
NAME SPECIMENS RIBS
RIBS/MM
HEIGHT
DIAMETER
H/D RATIO
IRREGULAR
thaumasius
Lauopa
mbalavuana
Tulmallla
infundibulum 156
42
44.1(36-55)
39
88.3(75-96)
91.2(74-103)
2.17(1.84-2.50) 2.72(2.30-3.32)
4.08(3.59-4.74) 1.28(1.12-1.51) 3.49(3.19-3.98)
1.86 3.24(2.81-3.66) 7.52(6.67-8.37)
4.88(4.17-5.37) 3.49(2.88-4.18) 5.68(4.97-6.86)
4.37(3.55-4.97) 3.69(3.07-4.12) 6.64(5.88-6.99)
0.801(0.752-0.842)
0.365(0.343-0.381)
0.430(0.422-0.437)
0.616(0.547-0.710)
0.556(0.495-0.606)
humbu.
thaum.
mbala.
inf un.
pilsb.
WHORLS
UMBILICUS
3 3/8(3 1/8-3 1/2) C L
4 1/8(4-4 3/8) 1.16(1.05-1.32)
4 1/2(4 1/4-4 3/4) 2.75(2.42-3.07)
3 3/4+(3 1/2-4 1/8) 1.04(0.86-1.32)
4+(3 3/4-4 3/8)
1.19(1.05-1.24)
D/U RATIO
S E D
2.99(2.82-3.15)
2.74(2.72-2.76)
5.47(4.47-7.15)
5.62(4.48-6.50)
APICAL
CORDS
12.0(11-13)
12.2(10-15)
22
MANY
REDUCED
MANY
REDUCED
thaum.
mbala.
SPIRE
ELEVATION
0.37(0.25-0.46)
DEPRESSED
0.15(0.07-0.23)
0.36(0.20-0.53)
0.47(0.33-0.66)
BODY WHORL
WIDTH
0.97(0.82-1.12)
0.75(0.66-0.84)
2.03(1.76-2.28)
1.94(1.74-2.07)
2.06(1.91-2.20)
SP/BWW
0.379(0.300-0.483)
0.069(0.037-0.100)
0.185(0.113-0.258)
0.223(0.159-0.308)
Free oviduct (UV) surmounted by a retractor muscle, swollen
above, tapering rapidly with muscular walls, then suddenly expand-
ing to form a thin-walled sac gradually tapering to spermathecal
junction. Spermatheca (S) with ovate head lying next to albumen
gland, shaft slender to base of uterine fold, lower portion rapidly
expanding to equal penis in width. Vagina (V) equal in diameter to
median free oviduct, very short, without prominent internal fea-
tures.
Free muscle system unusual only in having a very large muscle
from the free oviduct apex attached to columellar retractor just above
tentacular retractors. Right ommatophoral retractor passing
through penioviducal angle. Tentacular retractors unite with tail fan
just above junction of latter with buccal retractor.
Stomach occupying % whorl after reaching expanded size. Intes-
tinal loops highly compacted, hindgut typical.
(Based on FMNH 152378, 4 adults 5.9-6.1 mm. in diameter.)
Tuimalila infundibulus (Hombron & Jacquinot,
1841). Figure 78d-f.
Helix infundibulum Hombron & Jacquinot, 1841, Ann. Sci. Nat.
Zool., (2) 16, p. 64 Vavao, Tonga Islands; Hombron & Jac-
quinot, 1852, Voy. Pol. Sud, Astrolabe et Zelee, Atlas, pi. 6, figs.
25-28 Vavao, Tonga; Rousseau, 1854, Voy. Pol. Sud, Astrolabe
et Zelee, Atlas, 5, p. 20.
Patula (Patula) radicalis Mousson, 1871, J. de Conchyl., 19, pp.
12-13, pi. 3, fig. 3 Vavao, Tonga Islands.
Helix radicalis (Mousson), Pfeiffer, 1876, Monog. helic. viv., 7, p.
164.
Helix crebriflammis Pfeiffer, 1876, Monog. helic. viv., 7, p. 148
lists infundibulum Hombron & Jacquinot, 1841, as a question-
able synonym of this New Zealand species.
Charopa radicalis (Mousson), Tryon, 1886, Man. Conchol., (2) 2, p.
210.
Endodonta (Flammulina) radicalis (Mousson), Hedley, 1893,
Nautilus, 7 (3), p. 35.
Endodonta (Charopa) radicalis (Mousson), Pilsbry, 1893, Man.
Conchol., (2)9, p. 35.
Rhytida (Ouagapia) radicalis (Mousson), Mollendorff, 1903, Syst.
Conchyl. Cab., I, 12, b, p. 83, pi. 13, figs. 12-15.
Diagnosis. Shell very large, diameter 4.97-6.86 mm. (mean
5.68 mm.), with 3!^-4'/e rather loosely coiled whorls. Apex and spire
moderately and evenly elevated, last whorl descending much more
rapidly, spire protrusion a little less than l h body whorl width, H/D
ratio 0.547-0.710 (mean 0.616). Apical whorls worn in all examples
examined, occasionally fine traces of weak and crowded spiral cords
visible under oblique lighting. Postnuclear sculpture of high, thin,
sharply defined, strongly protractively sinuated radial ribs, 75-96
(mean 88.3) on the body whorl, whose interstices are 2-4 times their
width. Ribs/mm. 4.17-5.37 (mean 4.88). Microsculpture a lattice of
very fine radial and spiral riblets, 4-10 microradials between each
pair of major ribs. No secondary spiral cording. Umbilicus narrow,
V-shaped, regularly decoiling, contained 4.47-7.15 times (mean
5.47) in the diameter, margins rounded. Whorl contours and aper-
182
SOLEM: ENDODONTOID LAND SNAILS
DG
PP
FIG. 79. Anatomy of the Tongan Tuimalila pilsbryi, new species. Station T-22, Eua, Tonga. FMNH 152378: a, pallial region; b, pallial and
terminal genitalia; c, interior of penis, epiphallus and vas deferens; d, detail of major penial stimulator; e, talon and hermaphroditic duct. Scale
lines equal 1 mm. (SH).
tural inclination normal. Color with narrow, wavy reddish flammu-
lations, a tendency for broad circular supraperipheral and basal red-
dish zones evident in many individuals.
Tuimalila infundibulus (fig. 78d-f) is smaller, has
a less protruded spire, but a greater H/D ratio than
does T. pilsbryi (fig. 78a-c). The latter has less crowded
ribbing, and the apical whorls show a clear radial ele-
ment (fig. 2c, p. 11) that apparently is lacking in T.
infundibulus. No other Polynesian Charopidae even
approach the size of this species.
Description of radicahs. Shell very large, with 4 rather loosely
coiled whorls. Apex and spire moderately elevated, slightly rounded
above, body whorl descending more sharply, H/D ratio 0.634. Apical
whorls 1%, sculpture of extremely fine and crowded spiral ribs that
are barely visible except under strong lateral lighting. Postnuclear
whorls with irregular, high, rounded, strongly protractively sinuated
radial ribs, about 92 on the body whorl, whose interstices are 2-3
times their width. Microsculpture of very fine, crowded radial riblets,
crossed by slightly finer and more crowded spiral riblets. Sutures
deep, whorls shouldered above, strongly flattened laterally above
periphery, with evenly and gently rounded subperipheral margins.
Color light yellowish brown with very irregular radial reddish macu-
lations that coalesce to form a broad and a narrow band above the
periphery, a wide subperipheral plus a faint umbilical color band.
Umbilicus narrowly open, V-shaped, regularly decoiling, contained
5.55 times in the diameter. Aperture subcircular, strongly flattened
laterally above periphery, inclined almost 25 from the shell axis.
Height of lectotype 3.82 mm., diameter 6.02 mm.
Lectotype of radicalis. Tonga: Vavau. Collected
by Dr. E. Graeffe. Zoologisches Museum der Univer-
sitat Zurich.
Range. Vavau Island, Tonga.
Paratypes. Zurich, FMNH 116982.
Material. Vavau (20 specimens, RSM, AMS,
FMNH 46222, SMF 165340, BPBM 54361, BPBM
167432): Talau Id., 1 mile west (Station T-7) of Neiafu
at 400 ft. elevation in dense low forest (62 specimens,
SYSTEMATIC REVIEW
183
FMNH 152423); Mo'ungalafa, near Tu'anuku (Station
T-9), about 14 miles southwest of Neiafu at 700 ft. ele-
vation in heavy forest (2 specimens, FMNH 152207);
Muitoulo (Station T-10), about 15 miles west of Neiafu
at 500 ft. elevation in heavy forest (19 specimens,
FMNH 152290); Pangai Motu Id. (Station T-ll), 3
miles south of Neiafu at 200 ft. elevation (4 specimens,
FMNH 152357); Makave village (Station T-12), 2%
miles southeast of Neiafu at 200 ft. elevation (1 speci-
men, FMNH 152348); Longomapu (Station T-14), 16
miles southwest of Neiafu in heavy forest at 400 ft.
elevation (14 specimens, FMNH 152380); Leimatu'a
(Station T-17), 9 miles north of Neiafu at 500 ft. eleva-
tion (2 specimens, FMNH 152323). "Tongatapu" (error)
(22 specimens, Zurich, FMNH 116983). Erroneous data
(3 specimens, FMNH 46243, 1KB).
Remarks. The name Helix infundibulum gener-
ally has been overlooked in the literature since Pfeiffer
(1876) tentatively suggested it might be a synonym of
the New Zealand Flammulina crebriflammis (Pfeiffer).
Comparison of the type illustrations of infundibulum
and radicalis leaves no doubt that they depict the same
species. I could not locate type material of Helix infun-
dibulum.
Mollendorff s reference of this species to Ouagapia
supposedly was based on work by Semper showing this
species to have paryphantid dentition. I suspect that
this was caused by a misreading of Semper (1874, p.
136), who stated that O. gradata was a paryphantid
and that probably many of the other "Patula" from the
same area were also paryphantids.
Extensive collections on Tongatapu in 1966 by
Laurie Price failed to reveal any specimens of this
species, hence I consider the material in the Mousson
collection to be mislabeled. Unfortunately, all of the
material taken on Vavau was dead so that Tuimalila
infundibulus could not be dissected. Collections were
made during a severe drought, and no living material
was obtained ofSinployea vicaria paucicosta from some
of the same stations.
At Station T-7 one individual had the remains of
an egg capsule with an embryonic shell dried inside.
Despite the narrow umbilicus, evidently the eggs are
deposited within the umbilical opening. Although this
practice is common in the Endodontidae (Solem, 1976b,
pp. 26-30), this is the only Polynesian Charopidae that
showed evidence of this trait.
Genus Lagivala, new genus
Generally minute Charopidae with 3%-4% tightly coiled whorls.
Apex and spire depressed (demani), flat (davidi), to moderately ele-
vated, body whorl descending much more rapidly. Apical sculpture of
10-14 fine spiral cords. Postnuclear sculpture of prominent to very
fine (minusculus) radial ribs, microsculpture typical, no secondary
spiral cording. Umbilicus very widely open, cup-shaped, narrow only
in minusculus. Periphery usually strongly compressed laterally (ex-
cept microglyphis), umbilical margin rounded (microglyphis, minus-
culus} to strongly shouldered (macroglyphis) . Sutures channeled in
macroglyphis, microglyphis, and minusculus. Shell unicolored with-
out darker flammulations. Parietal wall with 2 or 3 large barriers
(only 1 in minusculus), extending posteriorly from 3 Ae of a whorl to
line of vision; where 2 parietals only, apparently a 3rd is sometimes
displaced onto columellar margin by partial detachment of parietal
wall. Palatal barriers 3 to 6, short, very deeply recessed crescents or
elongated lamellar blades situated near posterior margin of parie-
tals, in minusculus only a long single barrier lying partly on col-
umellar and partly on basal margin. Anatomy unknown.
Type species. Lagivala vivus, new species.
Lagivala and Vatusila present generally contrast-
ing patterns of structure (fig. 80-85). Lagivala has the
body whorl laterally compressed (except for micro-
glyphis and some demani); Vatusila has the body whorl
flattened above the periphery. Lagivala lacks secon-
dary spiral cording; Vatusila (except vaitupuensis) has
secondary spiral cording. Lagivala has deeply recessed,
short, and crescentic palatal barriers (except minus-
culus and some demani); Vatusila has fewer, much
longer, and less deeply recessed palatals. Lagivala has
very long and rather complexly twisted, high parietals;
Vatusila shows a strong tendency toward fusion and
reduction of the parietals. Lagivala has a very wide,
cup-shaped umbilicus (except minusculus); Vatusila a
U-shaped, narrower umbilicus (cup-shaped only in
eniwetokensis). Lagivala has a relatively low spire;
Vatusila, a high spire (except eniwetokensis). They
agree in major sculpture and rib spacing, but only be-
cause both taxa exhibit an unusually wide range of
types. Several Lagivala show a tendency toward chan-
neling of the sutures, but others show no evidence of
this change. Despite overlapping distributions and
lack of anatomical evidence, I have no hesitation in
considering them to be phyletically separate experi-
ments in apertural constriction.
From the above list of comparisons, it is evident
that two species are atypical. Vatusila eniwetokensis
has the flat spire and cup-shaped umbilicus of
Lagivala. Its possession of prominent secondary spiral
cording (fig. 86f), gradual descension of the upper
parietal (fig. 85e), rounded body whorl with flattening
restricted to above the periphery, shorter parietals,
and relatively narrow umbilicus (table XL) are char-
acters that I consider to be more significant than the
probably linked spire and umbilical shape changes.
Lagivala minusculus may be incorrectly placed here,
but insufficient data are available to propose generic
separation. Temporary classification within Lagivala
is a conservative and reasonable treatment pending
more information.
Similarities to other genera are few. Graeffedon
(figs. 86, 88) is much, much larger with short simple
parietals and narrower, much less deeply recessed
palatals that have abrupt anterior descension, a nar-
rower, V-shaped umbilicus, and prominent, darker
color flammulations. Other Melanesian and Polyne-
sian taxa lack any apertural barriers. Of the Microne-
sian genera, Palikirus (fig. 91d-f) has only a single
parietal barrier, a much narrower, V-shaped um-
bilicus, and much more widely spaced ribbing. Jokaj-
don (fig 92a-f) has very fine ribbing, many complicated
apertural barriers, and secondary spiral cording. Pal-
184
SOLEM: ENDODONTOID LAND SNAILS
davidi
(Ladd)
macroglyphis
(Rensch)
microglyphis
(Rensch)
TABLE XL. - RANGE OF VARIATION IN LAGIVALA.
"RHBER OF
SPECIMENS
1
2
1
82
5
RIBS/:*.
70 or 87
15.9 or 19.8
62.5(62-63)
11. Ii7(11.39-ll. 55)
121
214.9
62.5(51-71-)
11.16(9.86-12.1(6)
95.5(86-101)
17.9(ll.8-20.8)
demani
(Tapperone-
Canefri )
( Ladd )
vivus
minusculus
macroglyphis
(Rensch)
microglyphis
(Rensch)
demani
(Tapperone-
Canefri )
0.8k
0.63
0.95(0.89-1.02)
0.71(0.66-0.714)
0.85(0.76-0.95)
79.0(56-106)
DIAMETER
1.1.
1.75(1.72-1.77)
1.55
1.73(1.614-1.89)
1.63(1.55-1.714)
1.90(1.73-2.07)
13. 09(9. 85-19. It)
H/D RATIO
0.500
0. It83(0.l477-0. It 90)
O.l40k
0.553(0.539-0.580)
0.14314(0. Ii25-o.lt 57)
O.ltli8(0.l4l6-0.5l8)
davldl
vivus
davidi
vivus
minus,
mac ro .
micro.
WHORLS UMBILICUS 1/U KAIL) WICAL CORDS
3 1/2 0.51 2.62 UNKNOWN
4 0.68(0.67-0.69) 2.82(2.61-3.03) UNKNOWN
3 3/8 0.30 5.22 10
4 3/8-(4 1/8-4 5/8) 0.85(0.79-0.94) 2.03(2.00-2.08) 11-12
3 7/8(3 3/4-4) 0.65(0.59-0.72) 2.54(2.41-2.61) Circa 11
4 1/8+O 3/4-4 1/2) 0.72(0.58-0.82) 2.64(2.48-3.03) 12.1(10-14)
SPIRE
ELEVATION
FLAT
FLAT
0.05
0.07(0.04-0.10)
0.05
BODY WHORL
WIDTH
SP/BWW
0.51
0.59
0.42 0.118
0.54(0.53-0.59) 0.131(0.076-0.188) 2 1
0.54 0.091 2 1
PR C P
2 or 1 3 or 4
30 6
10 1
5
donanl FLAT OR DEPRESSED 0.61(0.58-0.67) FLAT OR DEPRESSED 2-3 0-1 T-l-1
line (figs. 94-96) has secondary spiral cording, looser
whorl coiling, and less deeply recessed palatals.
Known species of Lagivala are from Eastern In-
donesia and West Irian (L. demani); New Britain, Bis-
marck Archipelago (L. microglyphis and L. macro-
glyphis); Viti Levu, Fiji (L. vivus and L. minusculus);
and Funafuti, Ellice Islands (L. davidi). Except for the
New Britain collections in shore drift and some sets of
L. demani, only one or two examples are known of each
despite intensive collecting efforts. Additional species
will quite probably be discovered in intermediate areas
since few attempts have been made at sampling the
minute litter fauna.
The similarities of all Lagivala except L. minus-
culus are covered in the species accounts. Apparently,
partial detachment of the parietal wall has resulted in
the 3rd parietal becoming a columellar barrier in the
Bismarck species and some populations of L. demani.
Partial palatal barrier reduction in some examples of
L. microglyphis shows how the reduced number in L.
davidi could be derived. The channeled suture of L.
minusculus agrees with the Bismarck species, whereas
its peculiar columellar-palatal barrier could be derived
from the 3rd parietal or columellar by downward,
rather than upward, twisting of the anterior portion.
In previous publications (Solem, 1958b, 1959a) I
have used the genus Beilania Preston (1913, p. 433) for
L. demani and several other species. Despite intensive
search over 13 years, I have not been able to locate any
specimens of Beilania demani Preston, 1913, in
museum or private collections. Although the missing
types may yet appear, the probabilities are that they
will not be located in the immediate future. Preston's
description places the parietals as being subperipheral,
and the palatal wall is without barriers. In the absence
of any specimens, I am reluctant to use the name
Beilania for this group until the structures of the type
species are known in regard to spire elevation, form
and length of the parietals, actual rib count, and actual
absence of the palatal barriers. Possibly Lagivala and
Beilania are synonymous, but it is equally possible
that Beilania inopina is more closely related to the
Philippine-Indonesian species, Beilania philippinensis
(Semper, 1874) (see Solem, 1957, pp. 7-8, fig. 3). That
species has a narrower, V-shaped umbilicus, looser
whorl coiling, a single crescentic parietal with gradual
anterior descension, and three nodular palatals that
are near the lip margin and often on a heavy callus
much as in Palline notera and P. biakensis. The two
latter species (figs. 94-96) differ in having prominent
secondary spiral cording, tighter coiling, a much
thicker body whorl, narrower umbilicus, and very
differently shaped apertural barriers. Beilania philip-
pinensis, although not dissected, shows major con-
chological differences from Lagivala demani and the
more eastern species of Lagivala. Rather than propose
an additional generic name for the Philippine-
Indonesian species, I prefer to use Beilania for that
species and the genotype, pending availability of addi-
tional material and soft parts for dissection.
Lagivala is the Fijian nickname, meaning "the
long fellow," applied to the famous chief Qaraniqio or
Dakuwaqa of the Rewa District in Viti Levu, Fiji.
Lagivala davidi (Ladd, 1968).
Ptychodon species A, Ladd, 1958, J. Paleont., 32 (1), p. 189, pi. 30,
figs. 13-15166-170 ft. in deep boring at Funafuti, Ellice Is-
lands (Pleistocene or Recent).
Ptychodon davidi Ladd, 1968, J. Paleont., 42 (3), p. 857, fig. 1170
ft. in deep boring, Funafuti, Ellice Islands.
Diagnosis. A very small species with less than 3M> tightly
coiled whorls. Apex and spire flat, body whorl descending slightly,
H/D ratio about 0.500. Apical whorls IMs, sculpture of fine spiral
cords. Postnuclear whorls with prominent, rounded, almost verti-
cally sinuated radial ribs, about 70-87 on the body whorl. Micro-
sculpture of fine radial and finer spiral riblets. Umbilicus widely
open, cup-shaped, regularly decoiling, contained about 2.60 times in
the diameter, margins weakly shouldered. Sutures deep, whorls
strongly rounded above and on basal margin, compressed laterally,
with evenly rounded outer margin. Parietal barriers 2, agreeing in
SYSTEMATIC REVIEW
185
structure with the upper pair in L. vivus. Outer wall with 3 deeply
recessed palatals, short and crescentic, a 4th barrier located at baso-
columellar margin of adults that may be either a columellar or a
palatal barrier.
Lagivala vivus (fig. 80a-c) differs fromL. davidi in
having three parietals, six palatals, and fewer major
radial ribs. Lagivala minusculus (fig. 80d-f) has only a
single parietal and one peculiar palatal. Lagivala mac-
roglyphis and L. microglyphis (fig. 81a-f) have ele-
vated spires and more palatal barriers.
Holotype. Deep boring from Funafuti, Ellice Is-
lands, at 170 ft. depth. AMS F.52343.
Range. Fossil at Funafuti, Ellice Islands.
Material. Only the holotype and lost specimen
initially studied by Ladd (1958) are known.
Remarks. The juvenile holotype has three deeply
recessed palatals in the aperture, whereas the lost
example had an additional barrier at or near the baso-
columellar margin. I am uncertain whether this was a
columellar or palatal barrier. Differences fromL. vivus
are covered in the diagnosis.
Lagivala vivus, new species. Figure 80a-c.
Diagnosis. Shell very small, diameter 1.72-1.77 mm. (mean
1.75 mm.), with about 4 tightly coiled whorls. Apex and spire flat or
very slightly depressed below level of penultimate whorl, body whorl
descending moderately, H/D ratio 0.477-0.490 (mean 0.483). Apical
sculpture of fine and crowded spiral cords, worn off over most of
surface. Postnuclear whorls with high, rounded, sharply defined, al-
most vertically sinuated, rather crowded radial ribs, 62-63 (mean
62.5) on the body whorl, whose interstices are 2-4 times their width.
Ribs/mm. 11.3911.55 (mean 11.47). Microsculpture of fine radial
riblets, 5-8 between each pair of major ribs, crossed by extremely
fine and crowded spiral riblets. No secondary spiral cording present.
Umbilicus widely open, cup-shaped, regularly decoiling, contained
2.61-3.03 times (mean 2.82) in the diameter, margins slightly shoul-
dered, walls of umbilicus moderately flattened internally. Sutures
deep, whorls strongly rounded above and on basal margin, com-
pressed laterally with evenly rounded outer margin. Parietal bar-
riers 3, extending posteriorly more than Vt whorl: upper a high, thin,
bladelike lamella, slightly elevated and expanded above on posterior
eighth, with abrupt descension at anterior end after even elevation
from posterior portion; 2nd slightly higher and more broadly ex-
panded on posterior quarter, equal in height to 1st during middle
section, with sinuated descension over anterior 8th; 3rd located just
above parietal-columellar margin, much lower than 2nd, weakly ele-
vated and expanded above on posterior half, with gradual descension
to point just inside lip edge. Columellar wall without barriers.
Palatal barriers 6, deeply recessed, short and crescentic: lower at
baso-columellar margin, relatively low, visible only by tilting of
aperture; 2nd twice height of 1st, longer, slightly flattened on top,
with relatively sharp anterior descension; 3rd and 4th slightly
shorter and % height of 2nd with more gradual anterior descension;
5th supraperipheral, equal in height to 2nd, with more gradual an-
terior descension; 6th midway to palatal-parietal margin, a deeply
recessed, low, and lamellate tubercle.
Lagivala vivus is readily identified by possessing a
third parietal and six palatals. Lagivala davidi has
more ribs, only two parietals, and three palatals, but
obviously is closely related. The Bismarck L. macro-
glyphis and L. microglyphis (fig. 81a-f) have much
more elevated spires, two parietals, a prominent col-
umellar barrier, and only five palatals. Lagivala de-
mani from Indonesia and West Irian has only three or
four palatals, a depressed spire, and usually only two
parietals.
Description. Shell very small, with 4 tightly coiled whorls.
Apex and spire slightly depressed, body whorl descending moder-
ately, H/D ratio 0.477. Embryonic whorls almost 1%, traces of fine
spiral cording present, surface mostly eroded. Postnuclear whorls
with high, prominent, sharply defined, crowded radial ribs, 63 on the
body whorl, whose interstices are 2-4 times their width. Micro-
sculpture of fine radial riblets, 5-8 between each pair of major ribs,
crossed by barely visible and extremely crowded spiral riblets. Su-
tures deep, whorls strongly rounded above and on basal margin,
compressed laterally, with evenly rounded outer margin. Umbilicus
broadly open, cup-shaped, regularly decoiling, contained 2.61 times
in the diameter, margins shouldered, inside walls flattened. Color
uniform light reddish yellow, without distinct flammulations. Aper-
ture ovate, compressed laterally, with evenly rounded outer margin,
inclined about 5 from shell axis. Apertural barriers as in diagnosis
above. Height of holotype 0.86 mm., diameter 1.76 mm.
Holotype. Fiji: Viti Levu, Station 54, hillside at
Telenaua, 48.3 miles west of Suva by Belt Road at 60 ft.
elevation, back of Serua village. Collected in leaf
mould by Yoshio Kondo, Elwood Zimmerman, and C.
M. Cooke, Jr., on July 26, 1938. BPBM 178598.
Range. South coast of Viti Levu, Fiji.
Paratype. Viti Levu: Lami Ridge, west of Suva
(Station 47) on limestone rock (1 specimen, BPBM
178488); Nandarivatu (Station F-16), ridge slope at
2,500 ft. elevation (1 specimen, FMNH 168364).
Remarks. Only three dead specimens are known.
They agree in apertural features, which clearly distin-
guish them from other Lagivala. Attempts in 1962 to
collect additional specimens were unsuccessful, and in
1965 L. Price obtained one dead adult.
The name vivus is to contrast this species with its
nearest relative, Lagivala davidi (Ladd, 1968) from
Funafuti Pleistocene or Holocene deposits.
Lagivala minusculus, new species. Figure 80d-f.
Diagnosis. A very small species with narrow umbilicus,
slightly elevated spire, numerous and crowded ribs, 1 parietal, and 1
sinuately twisted palatal.
Other Lagivala have much more widely open um-
bilici, two or three large parietals and three to five
deeply recessed palatals. The singular palatal of L.
minusculus immediately separates it from any of the
Vatusila with reduced barrier number.
Description. Shell very small, with 3% tightly coiled whorls.
Apex and spire slightly and evenly elevated, body whorl descending
slightly more rapidly, spire protrusion about l k> body whorl width,
H/D ratio 0.404. Apical whorls l'/2, sculpture of 10 fine spiral cords,
whose interstices are about 3-5 times their width. Postnuclear
whorls with narrow, prominent, sharply defined, slightly sinuately
protractive radial ribs, 121 on the body whorl, whose interstices usu-
ally are less than twice their width. Ribs/mm. 24.9. Microsculpture of
very fine radial riblets, 2-4 between each pair of major ribs, crossed
by slightly finer and more crowded spiral riblets. Sutures distinctly
channeled after embryonic whorls, whorls strongly rounded above,
moderately compressed laterally and on basal margin. Umbilicus
relatively narrow, U-shaped, slightly and regularly decoiling, con-
tained 5.22 times in the diameter, margins rounded. Color light red-
dish yellow-horn, no flammulations. Aperture ovate, compressed lat-
erally and slightly on basal margin, inclined about 10 from shell
axis. Parietal wall with single medial, high, bladelike barrier,
slightly twisted upward posteriorly, weakly expanded above, extend-
abc
h
def
FIG. 80. a-c, Lagivala I'iuus, new species. Station 54, Telenaua, west of Suva, Viti Levu, Fiji. Holotype. BPBM 178598; d-f, Lagivala
minusculus, new species. Nangava Island, Viti Levu, Fiji. Holotype. BPBM 179255. Scale lines equal 1 mm. (MM).
186
SYSTEMATIC REVIEW
187
ing posteriorly beyond line of vision, without noticeable descension
until just before anterior end. Outer wall with a single moderately
elevated, bladelike barrier, posteriorly at line of vision on columellar
wall, anteriorly twisting down onto basal margin, reaching lip edge
with sharp anterior descension, buttressed on outer side by a thick,
tapering callus. Both barriers minutely serrated above. Height of
holotype 0.63 mm., diameter 1.55 mm.
Holotype. Fiji: Nangava Island, IVa miles south
of Viti Levu, near Namuka Island, at 210 ft. elevation.
Collected under dead hau leaves by Yoshio Kondo on
September 20, 1938. BPBM 179255.
Range. Nangava Island, south of Viti Levu, Fiji.
Remarks. Inclusion of this species in Lagivala is
based upon the channeled suture (also in L. macro-
glyphis andL. microglyphis), shape of the parietal bar-
rier (as inL. viuus andL. davidi), and general similar-
ity of the columellar-palatal barrier to the third
parietal in L. vivus (fig. 80b). The absence of any
palatal barriers, narrow umbilicus, much more
crowded and narrower ribbing, plus the peculiar ter-
mination of the columellar-palatal differ from the
Lagivala pattern. It is quite possible that L. minus-
culus represents a distinct lineage and should be gen-
erically separated. Without more specimens and mate-
rial for dissection I prefer to include it in Lagivala.
Lagivala macroglyphis (I. Rensch, 1937). Figure
81d-f.
Ptychodon macroglyphis I. Rensch, 1937, Arch. f. Naturgesch., n.
f., 6 (4), p. 592, fig. 36 right Karlei, Malkong, and Matong, New
Britain, Bismarck Archipelago.
Diagnosis. Shell minute, diameter 1.64-1.89 mm. (mean 1.73
mm.), with 4'/8-4 5 /8 rather tightly coiled whorls. Apex and early spire
flat or slightly depressed to barely elevated, last whorl descending
much more rapidly, spire protrusion slightly more than Vg body whorl
width, H/D ratio 0.539-0.580 (mean 0.553). Apical whorls 1%-!%,
sculpture of 11-12 fine spiral ribs whose interstices are about twice
their width. Postnuclear sculpture of large, high, vertically sinuated
radial ribs, 51 and 74 on 2 adult examples, whose interstices are 3-5
times their width. Ribs/mm. 9.86-12.46 (mean 11.16). Micro-
sculpture a lattice of very fine, almost coequal spiral and radial rib-
lets, 6- 10 between each pair of major ribs. Umbilicus broadly open,
cup-shaped, regularly decoiling, contained 2.00-2.08 times (mean
2.03) in the diameter, margins distinctly shouldered, inner walls
flattened, sutures strongly channeled with part of parietal wall de-
tached. Sutures channeled above, whorls strongly rounded above and
on basal margin with evenly rounded, strongly compressed laterally
on outer margin, umbilical margin shouldered. Color partly leached
from shell, a reddish yellow tint remaining. Aperture compressedly
ovate, strongly rounded above and below, markedly compressed lat-
erally, lying parallel to shell axis. Parietal barriers 2, extending
posteriorly almost to line of vision: upper a high, thin blade with
abrupt anterior descension, becoming thickened above on posterior
8th with slight downward twist or weakly bifid; lower parietal with
anterior 8th a high threadlike ridge, abruptly becoming equal in
height to upper parietal, greatly thickened, expanded and elevated
above on posterior quarter. Columellar wall with single bladelike
barrier situated just below parietal-columellar margin on detached
parietal wall, identical in form to 2nd parietal, but only half as high
and extending not as far anteriorly. Palatal wall with 5 thin, deeply
recessed, very short, crescentic ridges situated opposite posterior end
of parietal barriers: 1st palatal basal in position, a very small, short
crescent; 2nd a high crescentic barrier, weakly expanded above,
whose apex lies opposite columellar; 3rd slightly lower than 2nd,
situated subperipherally and directly opposite lower parietal; 4th
equal in size to 3rd, situated opposite upper parietal; 5th smaller
than 1st, a tiny short, low barrier near upper palatal margin, occa-
sionally absent. All barriers minutely serrated above.
The sympatric Lagivala microglyphis (fig. 81a-c)
differs from L. macroglyphis (fig. 81d-f) in having a
distinctly narrower umbilicus, lower H/D ratio, finer
and more crowded radial ribs, plus slightly smaller av-
erage size. Lagivala vivus from Fiji lacks the partly
detached parietal wall and thus has three parietals. It
also has six palatals and a slightly depressed apex and
spire.
Holotype. Karlei, Weiten Bucht, New Britain,
Bismarck Archipelago. Collected by Schneider. ZMB.
Range. Karlei, Malkong, and Matong, New Brit-
ain, Bismarck Archipelago.
Material. New Britain: Karlei (81 specimens,
ZMB, FMNH 146021); Malkong-Bach, Karlei (25
specimens, ZMB); Matong (1 specimen, ZMB).
Remarks. Lagivala macroglyphis has several
unusual features produced by partial separation of the
postnuclear whorls during growth. On the spire (fig.
81d) this is evidenced by deep channeling of the su-
tures that are partly filled by periostracal material in
fresh examples. In the umbilicus (fig. 81f) there is the
same partial detachment of the parietal wall seen in
the Marquesan Planudonta (Solem, 1976a, figs. 146,
149). The upper portion of each umbilical volution
lacks shell sculpture. This results from simple detach-
ment of the parietal callus. At the same time, the 3rd
parietal barrier of L. vivus has been lowered to occupy
the "columellar wall." It is, of course, completely
homologous with the parietal and is called a columellar
here only to facilitate identification by nonspecialists.
At its anterior end there is a pronounced tendency for a
distinct upward angling of the barrier to the functional
columellar-parietal margin.
Lagivala vivus (fig. 80a-c) lacks the whorl de-
tachment, has a narrower umbilicus, lower H/D ratio, a
flat spire, and an additional palatal lamella. In size
and sculpture it is quite similar.
Lagivala microglyphis (I. Rensch, 1937). Figure
81a-c.
Ptychodon microglyphis I. Rensch, 1937, Arch. f. Naturgesch., n. f.,
6 (4), p. 591, fig. 36 left Karlei, Weiten Bucht, New Britain,
Bismarck Archipelago; Solem, 1957, Fieldiana: Zoology, 42 (1),
p. 7.
Diagnosis. Shell minute, diameter 1.55-1.74 mm. (mean 1.63
mm.), with 3%-4 rather tightly coiled whorls. Apex and early spire
flat to barely protruding, last Vfe-% of body whorl descending a little
to much more rapidly, spire protrusion about 'Ai body whorl width,
H/D ratio 0.425-0.457 (mean 0.434). Apical whorls 1V4, sculpture of
11 prominent spiral cords in only unworn examples. Postnuclear
whorls with prominent, rounded, slightly protractively sinuated ra-
dial ribs, 86-101 (mean 95.5) on the body whorl, whose interstices
are 1-3 times their width. Microsculpture of very fine radial riblets,
5-8 between each pair of major ribs, crossed by coequal spiral riblets.
Umbilicus broadly open, cup-shaped, regularly decoiling, contained
2.41-2.61 times (mean 2.54) in the diameter, margins rounded. Su-
tures deep, very weakly channeled, whorls strongly rounded above
and on basal margin, nearly evenly rounded on outer margin with
very slight lateral flattening above and below periphery. Color red-
dish yellow-horn without darker markings. Aperture subovate,
FIG. 81. a-c, Lagivala microglyphis (Rensch). Karlei, New Britain, Bismarck Archipelago. Paratype. FMNH 146022; d-f, Lagivala mac-
roglyphis (Rensch). Karlei, New Britain, Bismarck Archipelago. Paratype. FMNH 146021. Scale line equals 1 mm. (SH).
188
SYSTEMATIC REVIEW
189
strongly rounded above and at basal margin, evenly rounded
elsewhere with slight lateral flattening, inclined about 20 from shell
axis. Parietal barriers 2, extending posteriorly to line of vision: upper
high and bladelike with very sharp anterior descension, slightly
twisted upward, posterior third to quarter varying from bifid to with
sharp downward deflection of greatly accentuated expanded portion;
lower parietal a more rounded ridge with very gradual descension
over anterior third, becoming higher, slightly bulbous on top, and
twisted upward on posterior third. Columellar wall with single,
bluntly rounded, moderately recessed, crescentic barrier extending
posteriorly about '/ whorl with gradual anterior descension across
top of columellar callus, slightly twisted upward over anterior third.
Palatal wall normally with 5 deeply recessed, short, and crescentic
barriers situated opposite posterior end of parietal barriers: lower
palatal basal in position, a short crescent, slightly bulbous above,
less than % height of next barrier; 2nd palatal high and crescentic,
slightly bulbous above, pointing toward upper margin of lower
parietal; 3rd palatal slightly smaller than 2nd, subperipheral, point-
ing between parietal barriers; 4th palatal equal in size to 2nd,
slightly supraperipheral in position, situated opposite edge of upper
parietal: 5th palatal equal in size to 1st, situated midway between
4th palatal and parietal-palatal margin. Occasionally only 3 palatals
are present, and an accessory trace may be present between various
lower palatal pairs. Expanded upper portions of each barrier with
minute crystalline barbs.
Lagiuala microglyphis has many more radial ribs,
a much lower H/D ratio, fewer whorls (table XL), and a
more rounded whorl contour than does the sympatric
L. macroglyphis (fig. 81d-f). In L. vivus (fig. SOa-c)
there are three parietals and no columellar, the latter
in both L. macroglyphis and L. microglyphis being a
descended third parietal.
Holotype. Karlei, Weiten Bucht, New Britain,
Bismarck Archipelago. Collected by Schneider. ZMB.
Range. Karlei, New Britain, Bismarck Ar-
chipelago.
Material. Karlei (5 specimens, ZMB, FMNH
146022).
Remarks. The five paratypes showed some bar-
rier variation. A small accessory crescentic barrier lies
between the second and third palatals of the largest
specimen; one adult had the first and fourth palatals
missing, with the second and third grossly thickened; a
juvenile lacked the first and fourth palatals, with no
thickening of the second and third palatals. The ten-
dency toward loss of two palatals provides a link to the
condition seen in L. davidi.
Suture channeling in L. microglyphis is greatly
reduced, and there is no indication of sculptural change
in the umbilicus caused by parietal wall descension,
despite the position of the columellar barrier. The
sculpture is much finer than in other Lagivala, except
for the unusual L. minusculus. Whether L. micro-
glyphis is an advanced or a generalized species cannot
be determined without additional material for study
and dissection.
Lagivala demani (Tapparone-Canefri, 1883).
Patula demani Tapparone-Canefri, 1883, Ann. Mus. Civ. Stor.
Nat., Geneva, 19, pp. 95-96, 300, pi. 2, figs. 13-15 Vokan, Aru
Islands; van Benthem Jutting, 1962, Ann. Mus. Civ. Stor. Nat.,
Genova, 73, p. 5.
Helix demani (Tapparone-Canefri), Tryon, 1887, Man. Conchol.,
(2) 3, pp. 26-27, pi. 4, figs. 39-41.
Nesophila demani (Tapparone-Canefri), Boettger, 1922, Abhl.
Senckenb. Naturf. Ges., 35 (4), pp. 373-374, pi. 21, fig. 5.
Beilania demani (Tapparone-Canefri), Solem. 1958, Arch. f. Mol-
lusk., 87 (1-3), p. 22; van Benthem Jutting, 1964, Nova Guinea,
Zool., 26, pp. 11-13 (partly) localities on Biak.
Ptychodon demani (Tapparone-Canefri), van Benthem Jutting,
1958, Nova Guinea, Zool., 9 (2), p. 327 Waima, Misool.
Diagnosis. Shell relatively large, diameter 1.73-2.07 mm.
(mean 1.90 mm.), with 3%-4'/4 tightly coiled whorls. Apex and spire
slightly and evenly elevated to moderately depressed below level of
penultimate whorl, often flat, body whorl descending much more
rapidly, spire protrusion at most 'A body whorl width, H/D ratio
0.416-0.518 (mean 0.448). Apical sculpture of 10-14 (mean 12.1)
very fine spiral cords, whose interstices are about 3 times their
width. Postnuclear whorls with prominent to very prominent, high,
sharply defined, usually slightly protractive radial ribs, 56-106
(mean 79.0) on the body whorl, whose interstices are 1V4-5 times
their width. Ribs/mm. 9.85-19.4 (mean 13.1). Microsculpture of fine
radial riblets, 3-13 between each pair of major ribs, crossed by
slightly finer and more crowded spiral riblets, no secondary spiral
cording. Umbilicus very widely open, V-shaped, regularly decoiling,
contained 2.48-3.03 times (mean 2.64) in the diameter, margins
rounded. Sutures deep, whorls strongly rounded above and on baso-
columellar margin, outer margin varying from markedly compressed
laterally to almost evenly rounded with slight lateral compression
above periphery. Aperture subcircular to compressedly ovate, in-
clined about 5-10 from shell axis. Parietal barriers normally 2
(only one known population with 3). high, extending posteriorly
3 fiB-Vt whorl: upper very high and slender with abrupt anterior de-
scension, posterior quarter often slightly more elevated with a
downward twist to elevated portion; 2nd usually equal in height on
crescentic posterior 3rd, which may be slightly twisted upward, mid-
dle 3rd slightly lower and less expanded or descending, with gradual
descension over anterior 3rd until just before termination slightly in
front of upper parietal; 3rd, when present, about '/2-% height of 2nd,
just above parietal-columellar margin. Columellar wall without any
barrier (Biak and West Irian), with a very small and deeply recessed,
ridgelike trace (Biak), a medium-size barrier (Ambon and Aru Is-
lands), or with barrier almost equal in size to 2nd parietal (Timor).
Palatal barriers 3 or 4, rarely absent, deeply recessed, variable in
size and position, crescentic and short to elongated lamellar ridges:
1st through 3rd on basal to outer lip, with 3rd just above periphery
and opposite upper parietal, sometimes shifted lower on outer wall;
4th, when present, midway between 3rd palatal and palatal-parietal
margin.
Lagivala demani differs from the Fijian L. vivus
and the two Bismarck species in its reduced number of
palatal barriers, generally lower H/D ratio, and
slightly greater size. Lagivala macroglyphis is much
more widely umbilicated, has a distinctly protruded
spire, and is very strongly compressed laterally on the
whorl margin. Lagivala microglyphis has finer radial
ribbing, is smaller, the body whorl has only slight lat-
eral compression, and the aperture is inclined about
20 from the shell axis. Lagivala vivus always has a 3rd
parietal and 6 palatals.
Description. Shell minute, with 3% moderately tightly coiled
whorls. Apex and spire slightly emergent, body whorl descending
only a little more rapidly, H/D ratio 0.454. Surface of shell heavily
encrusted with dirt and fungal growth so that sculptural details are
obscured. Size and position of major ribs as in "Diagnosis" cited
above. Umbilicus broadly V-shaped, widely open, regularly decoil-
ing, contained 2.54 times in the diameter. Apertural lip badly bro-
ken, obscuring shape of aperture, sutures filled with extraneous mat-
ter. Aperture showing only anterior edges of the 2 parietal barriers,
190
SOLEM: ENDODONTOID LAND SNAILS
otherwise choked with dirt and fungal hyphae. Height of lectotype
0.74 mm., diameter 1.63 mm.
Lectotype. Aru Islands: Wokan. Collected by O.
Beccari. MSNG.
Range. Ambon, Timor, Wokan in the Aru Is-
lands, Misool, Biak off West Irian, and coastal areas of
West Irian. Possibly Celebes.
Parades. MSNG, SMF 4678.
Material. Ambon (1 specimen, ZMA collected Oc-
tober 11-13, 1949, by M. A. Lieftinck). Timor: Araki-
Vlakte (5 specimens, FMNH 150775, ZMA collected
April 12, 1950, by B. Polak). Aru Islands: Wokan (5
specimens, MSNG, SMF 4678). Misool: Waima (6
specimens, FMNH 150774, ZMA collected September
13-14, 1948 by M. A. Lieftinck). Biak: Kombai (00 46',
135 58') (12 specimens, ZMA, RNHL); Korim (00 55',
136 04') (10 specimens, FMNH 150776, RNHL); Sta-
tion G-65, Saba (1 specimen, RNHL); Station G-34,
Owi, South Biak (1 specimen, RNHL); Station G-30,
Bosnik (1 specimen, RNHL). West Irian: Ugar, Kokas
District (1 specimen, RNHL collected June 6, 1953);
Sarmi (01 52', 138 44') (2 specimens, ZMA collected
December, 1962, by G. J. M. Gerrits).
Remarks. In the type lot of Lagivala demani
there were five examples of this species and a single
Discocharopa aperta (Mollendorff). Tapparone-
Canefri's original figures show an extremely widely
umbilicated shell without trace of apertural barriers.
Although the Lagivala were all juvenile and had the
aperture choked with dirt so that only the tips of the
parietals were visible, I suspect that, despite no men-
tion of the barriers, the description was based on the
Lagivala and the figures on the Discocharopa. Boettger
(1922) illustrated one specimen from the type lot with
the parietals and depressed spire. I prefer to use the
name in this sense rather than to adopt the Dis-
cocharopa as type, since its reduced sculpture and ele-
vated spire agrees more with the type figures.
Tapparone-Canefri (1883, p. 300) indicated in the ex-
planation of the plates that the sculpture was omitted.
The very fine ribs of the Discocharopa could have been
overlooked much more easily than the coarse ribbing of
the Lagivala. Recently, van Benthem Jutting (1962, p.
5) indicated that the lectotype was the "shell figures pi.
2, fig. 13-15." There is little agreement between the
lectotype and the figures.
Unfortunately, most available material was
juvenile. Only 12 of the Biak examples from Kombai,
Korim, and Saba plus one shell from Misool showed
adult shell features. Because of the barrier variation,
changes in spire elevation and ribbing and variation
data for several sets of adults and juveniles are sum-
marized in Table XLI. Size differences should be ig-
nored, because the sets are not comparable in age com-
position. Since umbilical decoiling is regular in this
species and the spire is normally flat or depressed, both
D/U and H/D ratios would be little affected by the
change from juvenile to adult growth. The very slightly
increased H/D ratio for the Timor shells is attributable
to the slightly elevated spire in that population. Shells
from Misool are more narrowly umbilicated, whereas
those from Korim, Biak, have a lower H/D ratio, possi-
bly as a correlative with the more depressed apex and
spire of that population. Rib counts vary widely, but
the more critical factor of rib spacing shows agreement
between the samples. Timor shells do have more prom-
inent ribs, but there is only a slight decrease in rib
spacing that is not statistically significant. Misool
shells have narrower and thus more numerous and
slightly more crowded major ribs (mean ribs/mm.
13.83) than other sets. Measured differences are thus
minor.
Changes in spire protrusion and apertural barriers
are much more significant. In the types from Wokan,
Aru Islands, the spire is barely elevated, the columel-
lar barrier is high but deeply recessed, and the three
palatals are situated in normal position, deeply re-
cessed, short and high. Juveniles from Timor in addi-
tion to the enlarged radial ribbing have the spire and
apex distinctly elevated, the columellar barrier is half
to two-thirds as high as the second parietal and ex-
tends to the top of the columellar callus with gradual
anterior descension, plus four palatals. The lower three
are high and crescentic, with the first at the baso-lower
palatal margin, the second pointing toward the lower
parietal top edge, and the third opposite and nearly
touching the upper parietal. A much smaller and lower
fourth palatal lies one-third of the way between the
third palatal and the palatal-parietal margin. One in-
dividual has an accessory cordlike trace at the baso-
columellar margin, more deeply recessed than the first
palatal and much shorter. All specimens from Biak
have a flat or noticeably depressed spire, the columel-
lar reduced to a threadlike or ridged lamellar trace
visible only by extreme tilting of the aperture, and the
three palatals longer, much lower, with the second
situated distinctly above the second parietal level, and
the third sometimes split into two parts. The Misool
examples have the spire flat or very slightly depressed,
no trace of any columellar barrier, and the position of
the palatals altered so that the first palatal lies at the
baso-columellar margin, the second at the baso-lower
palatal margin, and the third pointing between the
first and second parietals. In size these palatals are
intermediate between those of the Biak and Timor
examples. Their downward shift in position probably
reflects adjustments to the columellar barrier loss and
serves to restore balanced narrowing of the aperture.
The broken and worn individual from Ambon has a
medium-sized columellar barrier, the palatals shifted
as in the Misool shells, and the spire very slightly ele-
vated. Two shells from Sarmi, West Irian, have a third
barrier on the lower edge of the parietal wall and equal
in height posteriorly to the second parietal. Its anterior
descension is very gradual and terminates behind the
end of the first parietal. There is no columellar barrier
as such. One shell has the typical short and crescentic,
deeply recessed three palatals, the other lacks any
SYSTEMATIC REVIEW
TABLE XLI. - LOCAL VARIATION IN LAGIVALA DEMANI .
191
NUMBER OF
SPECIMENS
Sarmi, W. Irian 2
ZMA
Kombai, Blak 5
ZMA, RNHL
Korlm, Biak 7
RNHL
Saba, Biak 1
RNHL
Waima, Misool 5
ZMA
Araki-Vlakte .Timor 4
ZMA
Wokan, Aru 5
SMF 4678.MSNG
Ambon 1
ZMA
RIBS
RIBS /MM.
65.510.53 13.01.32
HEIGHT
0.76+0.049
(55-76)
(11.68-14.31)
(0.71-0.81)
DIAMETER
1.600.099
(1.50-1.69)
75.54.01 13.10.66 0.860.032 1.86+0.080
(56-106) (9.85-19.39) (0.76-0.93) (1.66-2.06)
70.1+3.37 12.30.47 0.7910.017 1.82+0.042
(56-85) (9.88-13.87) (0.72-0.86) (1.63-1.96)
57.7+0.33
0.95
2.07
71.8+6.65 13.8+0.41 0.75+0.044 1.610.101
(56-87) (12.92-14.85) (0.66-0.90) (1.38-1.97)
54.52.53 12.70.54 0.660.017 1.37+0.028
(47-58) (11.33-13.68) (0.63-0.71) (1.32-1.45)
13.0+0.61
(57-58) (12.31-14.20)
50 10.2
0.68+0.022 1.45+0.056
(0.64-0.74) (1.30-1.63)
0.74
1.56
H/D RATIO
Sarmi 0.475+0.0014
(0.473-0.476)
Kombai 0.4650.0134
(0.443-0.518)
Korim 0.4350.0040
(0.416-0.448)
Saba
0.460
WHORLS
4 1/8
4 1/4(4-4 1/2)
4(3 1/2-4 1/4)
4 1/2
3 l/2-(3 1/8-3 3/4)
UMBILICUS
D/U RATIO
Waima 0.4630.0118
(0.455-0.476)
Araki 0.4850.0100
(0.475-0.488)
Wokan 0.468+0.0099 3 l/2-(3 1/4-3 7/8)
(0.438-0.493)
0.660.033
(0.63-0.69)
2.410.030
(2.39-2.45)
0.69+0.046
(0.58-0.81)
2.71+0.086
(2.55-3.03)
0.71+0.020
(0.64-0.79)
2.55+0.021
(2.48-2.63)
Ambon
0.473
0.82
0.540.040
(0.41-0.66)
0.51+0.019
(0.46-0.54)
0.5510.031
(0.48-0.64)
0.63
2.52
2.99+0,
(2
2.66+0,
(2,
2.670
(2
2.50
,100
,81-3.36)
,076
,49-2.86)
,054
,53-2.78)
trace of palatals. Whorl contours vary with the ribbing,
the Sarmi and Misool shells having the body whorl
markedly compressed laterally, the Biak shells show-
ing only slight lateral compression, and the Timor
shells having a subcircular body whorl contour with
some lateral compression above the periphery. Reduc-
tion of the columellar barrier correlates with increased
lateral compression of the body whorl.
Recognizably different populations exist, and the
pattern of variation in many Wallacean animal groups
strongly suggests subspeciation has occurred. With
only single collections of mainly juvenile individuals
from widely scattered areas I prefer not to give
nomenclatural recognition to these populations. Varia-
tional data summarized above and in Table XLI should
enable interpretation of variation in additional popula-
tions but are not sufficient for description of subspecific
units.
The wide range of this species, extending as far
west as Timor, Ambon, and Misool, makes it quite
plausible that Beilania inopina Preston, 1913, from
Beilan-Beilan Island north of Obi is a member of this
complex. Reasons for not including it at this time have
been given above (p. 184).
A single worn and broken shell from Tjamba Pass,
near Makassar, South Celebes, was located in a set of
Discocharopa aperta. It may be an undescribed species
of Lagivala or represent a population of L. dernani in
which the palatals are very close to the lip.
Genus Vatusila, new genus
Shell relatively small to very small, with 3'/2-5 normally to
rather tightly coiled whorls. Apex and spire flat (eniwetokensis) to
rather strongly elevated (nayauana), body whorl usually descending
much more rapidly. Apical sculpture of 8-12 fine, but prominent
spiral cords. Postnuclear sculpture of crowded to widely spaced radial
ribs, strong secondary spiral cording in all but niueana (weakened)
and vaitupuensis (absent). Umbilicus rather widely open, narrowed
in kondoi. Periphery rounded, whorls slightly to strongly flattened
laterally above periphery. Color flammulations, when present, not
zigzagged, following lines of growth. Parietal wall with 2 barriers, 1
192
SOLEM: ENDODONTOID LAND SNAILS
bifurcated barrier, or greatly reduced single barrier. Columellar wall
with or without recessed, low ridge. Palatal barriers varying from
0-3, broadly expanded (tongensis) down to a low, broad remnant
callus (niueana). Anatomy mostly unknown, penis in tongensis
differing from Sinployea only in accentuation of semicircular pilas-
ters forming vergic papilla.
Type species. Vatusila tongensis, new species.
Despite the rather widely different appearance of
the sculpture and large size range, I have no hesitation
in considering these species congeneric. The two small
species, V. kondoi (fig. 82a-c) and V. nayauana (fig.
82d-f) from the Lau Group, and V. eniwetokensis (fig.
85e-f) from the Miocene of Eniwetok, agree in shell
microsculpture, although not in parietal barrier size or
character of major ribbing. The larger V. tongensis (fig.
83a-c) from Eua has the microsculpture of this group
and agrees in parietal barrier shape with the fossil.
Comparison of the figures shows that a callus connect-
ing the second and third palatals, elaboration of the
first palatal, and loss of the columellar from V. kondoi
would duplicate the outer lip barriers in V. tongensis.
Although no direct linear relationship between these
species should be implied, all the above species could be
derived from a common model. Vatusila vaitupuensis
(fig. 85a-d) and V. niueana (fig. 83d- f) would represent
successive stages in barrier reduction from a V.
tongensis barrier stage, although they agree more with
V. eniwetokensis in major rib spacing. They differ in
having modified secondary spiral sculpture or none
(vaitupuensis).
It is significant that those with the widely spaced
major sculpture, V. tongensis, V. kondoi, and V.
nayauana, are from a central area, whereas those with
more average sculpture, V. niueana, V. vaitupuensis,
and V. eniwetokensis, have a fringing distribution
around this core. I suspect that more species of this
group will be discovered in Fiji and Tonga when addi-
tional collections become available.
Comparisons with other genera are unsatisfactory
because so little anatomical data were available. The
penis of Vatusila tongensis (fig. 84a-b) represents a
rather minor change from the pattern seen in Sin-
ployea. Without more complete material for study, the
closeness of this relationship will remain uncertain.
The small species with heavier barriers show consider-
able sculptural modification from the pattern of
Sinployea, whereas one of the larger species, V.
niueana, can be mistaken for that genus very easily.
Progressive reduction in aperture barriers with in-
creased size is common in the Endodontidae, but this
cannot be postulated for Vatusila, because the sizes of
V. tongensis, V. vaitupuensis, and V. niueana are es-
sentially identical. Species ofLagivala have much sim-
pler structure to the parietal barriers; their palatals
are short, simple, and very deeply recessed. They differ
in both microsculpture and whorl contours and seem to
be a group paralleling the structures of Vatusila, but of
different origin. Microcharopa (fig. 33a-c) is smaller,
has greatly modified apical shell sculpture, and lacks
apertural barriers. Graeffedon (figs. 86, 88) is much,
much larger and has quite different palatal barriers
(although the parietals have a similar structure to
those found in V. eniwetokensis) and a very different
anatomy (fig. 87a-d).
Vatusila is named after the Fijian tribe (located at
the headwaters of the Sigatoka River) that killed and
ate Rev. Thomas Baker, a Wesleyan missionary, on
July 21, 1867.
Vatusila kondoi, new species. Figure 82a-c.
Diagnosis. Shell very small, diameter 1.68-1.84 mm. (mean
1.75 mm.), with 3%-3% normally coiled whorls. Apex and spire mod-
erately elevated, slightly rounded above, last whorl descending much
more rapidly, spire protrusion about Vs body whorl width, H/D ratio
0.500-0.610 (mean 0.552). Apical sculpture of 9-11 (mean 10.3)
rather fine spiral cords. Postnuclear whorls with high, prominent,
very widely spaced, strongly protractively sinuated radial ribs,
31-48 (mean 39.0) on the body whorl, whose interstices are 5-10
times their width. Ribs/mm. 5.51-8.06 (mean 7.04). Microsculpture
of barely visible radial and spiral riblets, almost completely obscured
by prominent and crowded secondary spiral cording. Umbilicus nar-
row, U-shaped, slightly and regularly decoiling, contained 4.12-5.40
times (mean 4.72) in the diameter, margins rounded. Sutures deep,
whorls strongly rounded above, flattened laterally with a weak su-
praperipheral sulcus, basal margin evenly rounded and slightly
compressed. Aperture ovate, inclined about 15 from shell axis.
Parietal wall with single high barrier, extending Vn- 3 he of a whorl
posteriorly, crescentic in outline with gradual anterior and posterior
descension, usually flattened above on middle third, varying from
slightly thickened medially to broadly bifid, with superior arm of
barrier twisted upward. Columellar wall with low, broadly rounded,
cordlike ridge, lying parallel to plane of coiling, reaching top of col-
umellar callus, occasionally (5.6%) absent. Palatal barriers usually
3, sometimes (11.1%) only 2, short, moderately to deeply recessed
within aperture: lower basal in position, crescentic, broadly rounded
above; 2nd much lower and slightly shorter; 3rd slightly sup-
raperipheral, longer, narrower, more deeply recessed bladelike bar-
rier with a very gradual anterior descension. Columellar and lower
palatal walls with a rather thick callus.
Vatusila kondoi (fig. 82a-c) is smaller and has
much smaller apertural barriers than the Eua Island
V. tongensis (fig. 83a-c). Vatusila nayauana (fig.
82d-f) has a distinctly higher spire, more crowded ra-
dial ribbing, a wider umbilicus (table XLII), and re-
duced apertural barriers. Vatusila eniwetokensis (fig.
85e-f) has much more numerous and crowded radial
ribs, two parietals, and a wider umbilicus.
Description. Shell very small, with 3'/2 moderately loosely
coiled whorls. Apex and spire moderately elevated, rounded above,
last whorl descending rapidly, H/D ratio 0.558. Apical whorls 1%,
sculpture of 11 relatively prominent, narrow spiral ribs, partly
eroded. Postnuclear whorls with widely spaced, protractively
sinuated, narrow, rounded radial ribs, 34 on the body whorl, whose
interstices are 5- 10 times their width. Microsculpture of very fine,
close-set radial ribs, crossing quite strong, relatively prominent sec-
ondary spiral cords. Microspiral sculpture barely visible. Major ribs
beaded on top by crossing of spiral cords. Sutures moderately deep,
whorls shouldered above, flattened laterally above periphery with
slight basal flattening, a weak supraperipheral sulcus present. Um-
bilicus narrowly U-shaped, regularly decoiling, slightly constricted
by expansion of basal lip, contained 4.72 times in the diameter. Color
uniform light yellow-brown. Aperture ovate, strongly flattened
above periphery, slightly so on base, inclined about 15 from the shell
axis. Parietal wall with a single high barrier, extending about 3 Ae of
a whorl, gradually descending anteriorly and posteriorly, broadly
expanded above and bifid in center. Columellar barrier a broad,
rounded, moderately prominent ridge, lying parallel to plane of coil-
a
FIG. 82. a-c, Vatusila kondoi, new species. Station 101, Nayau, Lau, Fiji. Holotype. BPBM 180259; d-f, Vatusila nayauana, new species.
Station 43, Nayau, Lau, Fiji. Holotype. BPBM 167232. Scale lines equal 1 mm. (MM).
193
194
SOLEM: ENDODONTOID LAND SNAILS
TABLE XLII. - RANGE OF VARIATION IN VATUSILA.
NUMBER OF
NAME SPECIMENS RIBS
RIBS /MM.
DIAMETER
H/D RATIO
kondoi
56
39.
0(31-48)
7
,04(5.51-8.06)
0.
97(0.86-1
,05)
navauana
42
51.
0(48-54)
9
,48(9.11-9.85)
0.
95(0.87-1
.02)
eniwe tokens is
1
89
15
,1
0.
86
(Ladd)
tongensis
20
61
,0(52-72)
7
.66(6.89-8.71)
1,
36(1.13-1
.72)
vaitupuensis
2
115
14
.27
1,
25
niueana
1
89
11
.60
1.
27
1.75(1.68-1.84) 0.552(0.500-0.610)
1.65(1.58-1.74) 0.573(0.552-0.608)
1.88 0.456
2.53(2.32-2.75) 0.538(0.472-0.650)
2.57 0.487
2.44 0.500
kondoi
WHORLS
3 l/2+(3 3/8-3 7/8)
4-(3 5/8-4 1/8)
3 1/2
4 5/8-(4 1/4-5)
4 1/2+
It 1/4
nayau.
eniwe.
tonge.
vai tu.
niuea.
APICAL
UMBILICUS D/U RATIO CORDS
0.37(0.35-0.43) 4.72(4.12-5.40) 10.3(9-11)
0.46(0.43-0.48) 3.61(3.43-3.69) 10.1(9-11)
0.53 3.56 12
0.76(0.66-0.89) 3.31(3.00-4.00) 10.4(8-12)
0.89 2.89 8
0.66 3.70 10
SPIRE
BODY WHORL
ELEVATION
UinTH
SP/BW
PR C
p
kondoi
.10
0.
56
0,
177
I 1 0-1
2-3
navau.
17(0.13-0.23)
0.
53(0.48-0.56)
0.310(0.260-0.420)
1 0-1
0-1
eniwe.
FLAT
0.
61
FLAT
2 ?
7
tonge.
.20(0.16-0.30)
0.
76(0.63-0.82)
0,
270(0.200-0.375)
I 1
2 2
vaitu.
0.13
0,
79
,167
1
2
niuea.
.13
0.
76
.174
1
Callu:
(Reduced)
1. Bifid
2. Enlarged laterally
ing, reaching top of columellar callus. Palatal barriers 3: lower
reaching edge of moderate apertural callus, short, >Ae whorl in
length, a high, broadly rounded ridge; middle palatal slightly more
deeply recessed, lower, narrower, an equally short threadlike ridge;
upper slightly supraperipheral in position, a broad V-shaped ridge,
short, with gradual anterior descension about as deeply recessed as
middle palatal. All barriers minutely serrated above and on sides.
Height of holotype 0.95 mm., diameter 1.71 mm.
Holotype. Fiji: Lau Group, Nayau, Station 101, %
mile north of Liku Village at base of a limestone cliff,
100 yd. inland at 30-500 ft. elevation. Collected in leaf
litter by Yoshio Kondo on August 22, 1938. BPBM
180259.
Range. Nayau, Lau Group, Fiji.
Paratypes. Lau Group: Nayau, % mile north of
Liku Village (Station 101) at base of a limestone cliff,
100 yd. inland at 30-500 ft. elevation (56 specimens,
BPBM 180259).
Remarks. The parietal barrier in Vatusila kon-
doi varies from a single lamellate ridge to a markedly
bifid barrier as in the type (fig. 82b). The columellar
barrier is often reduced in size, occasionally absent,
and the middle palatal may be absent. These varia-
tions are relatively minor and do not bridge the gap
between the barriers of V. kondoi and V. nayauana.
Differences in proportions (table XLII) are equally
large, and I have no hesitation in placing them as dis-
tinct species.
Great pleasure is taken in naming this species
after its collector, Dr. Yoshio Kondo.
Vatusila nayauana, new species. Figure 82d-f.
Diagnosis. Shell very small, diameter 1.58-1.74 mm. (mean
1.65 mm.), with 3%-4'/8 rather tightly coiled whorls. Apex and spire
rather strongly elevated, sometimes rounded above, last whorl de-
scending distinctly more rapidly, spire protrusion about Va body
whorl width, H/D ratio 0.552-0.608 (mean 0.573). Apical sculpture of
9-11 (mean 10.1) fine spiral cords, whose interstices are about 3
times their width. Postnuclear whorls with narrow, prominent,
rounded, protractively sinuated radial ribs, 48-54 (mean 51.0) on the
body whorl, whose interstices are 3-7 times their width. Ribs/mm.
9.11-9.85 (mean 9.48). Microsculpture of extremely fine radial and
spiral riblets, obscured by much more prominent, rather crowded
secondary spiral cords. Umbilicus rather widely open, broadly U-
shaped, last whorl decoiling more rapidly, contained 3.43-3.69 times
SYSTEMATIC REVIEW
195
(mean 3.61) in the diameter, margins rounded. Sutures deep, whorls
strongly rounded above, slightly flattened laterally above periphery,
with evenly rounded lower margins. Aperture ovate, inclined about
10 from shell axis. Parietal barrier a short, high, crescentic blade,
extending posteriorly about Vs whorl, expanded and serrated above,
with gradual anterior and posterior descension, medial in position.
Columellar wall with (60%) or without (40%) a low, rounded cordlike
ridge that barely reaches top of columellar callus. Palatal wall with
(60%) or without (40%) a single, bladelike barrier, relatively low,
strongly expanded and serrated above, with gradual anterior descen-
sion, moderately recessed, situated on middle of lower palatal mar-
gin.
Vatusila nayauana (fig. 82d-f) differs from V.
kondoi (fig. 82a-c) in its fewer and smaller apertural
barriers, much wider umbilicus, more protruded spire,
more crowded radial ribs, and greater H/D ratio (table
XLII). Vatusila eniwetokensis (fig. 85e-f) has two
parietals, much more numerous and crowded ribbing,
plus a flat spire. Vatusila tongensis (fig. 83a-c) is much
larger and has grossly developed apertural barriers.
Description. Shell very small, with slightly over 4 rather
tightly coiled whorls. Apex and spire moderately and evenly ele-
vated, last whorl descending more rapidly, H/D ratio 0.608. Apical
whorls 1%, sculpture of 11 narrow spiral ribs, whose interstices are
about 3 times their width. Postnuclear whorls with protractively
sinuated, widely spaced, prominent, rounded radial ribs, 48 on the
body whorl, whose interstices are 3-5 times their width, becoming
more crowded on the last half of body whorl. Microsculpture of very
fine and crowded, inconspicuous radial riblets and lower, much more
crowded spiral riblets, crossing stronger, rather widely spaced spiral
cords. Major ribs finely beaded above by crossing of spiral cords.
Sutures relatively deep, whorls strongly rounded above, slightly flat-
tened laterally above periphery. Umbilicus open, broadly U-shaped,
last whorl decoiling more rapidly, contained 3.64 times in the diame-
ter. Aperture circular, inclined about 10 from the shell axis with a
moderately prominent basal callus. Parietal wall with a short bar-
rier extending about Vs whorl, broadly rounded and ridgelike. Col-
umellar wall with a small, deeply recessed, cordlike tubercle. Palatal
wall with a subperipheral, broadly rounded ridgelike barrier, mod-
erately recessed, expanded above, extending about *Ae of a whorl.
Major palatal and parietal minutely serrated above and on sides.
Height of holotype 1.02 mm., diameter 1.68 mm.
Holotype. Fiji: Lau Group, Nayau, northwest
coast, Station 43, Nauko, hillside Vfe mile inland at 250
ft. elevation. Collected dead at the base of a limestone
cliff by H. S. Ladd on August 13, 1934. BPBM 167232.
Range. Nayau, Lau Group, Fiji.
Paratypes. Lau Group; Nauko, hillside, Vz mile
inland (Station 43) at 250 ft. elevation (42 specimens,
BPBM 167232).
Remarks. Although Vatusila nayauana can be
derived from V. kondoi by a combination of barrier
reduction, greater spire protrusion, rib crowding, and
umbilical widening, these changes are not correlated.
Hence specific-level recognition is warranted.
The exact relationship of the type localities is un-
certain. Vatusila nayauana was taken on the north-
west coast of Nayau, either at or on the way to Ladd's
geological station L502-L504 (Ladd & Hoffmeister,
1945, pp. 86, 190). Vatusila kondoi was taken "three-
quarters mile north of Liku Village." The latter is lo-
cated approximately in the middle of the southwest
coast of the island. Although this locality is 100 yd.
inland, Ladd's station was cited as Vfe mile inland.
Hence, regardless of the relative shore proximity of the
sites, they are separated at a minimum by about 700-
800 yd. Thus the snails are isolated colonies. Only five
examples were adult. Most of the rest had the aperture
clogged with debris. Hence the proportion of specimens
that lack the columellar and palatal barriers may be
altered considerably when new collections are avail-
able.
Vatusila eniwetokensis (Ladd, 1958). Figure 85e-f.
Ptychodon eniwetokensis Ladd, 1958, J. Paleont., 32 (1), p. 190, pi.
30, figs. 9-12 Drill hole K-1B at 820-831 ft. depth, Engebi
Islet, Eniwetok Atoll, Marshall Island (Miocene g).
Description. Shell very small, probably with 4 whorls when
complete (only 3'/2 remaining). Apex barely emergent, spire flat, last
whorl descending more rapidly, H/D ratio of remnant 0.456. Apical
whorls 1%, sculpture of 12 fine spiral ribs. Lower whorls with
crowded, prominently narrow, protractively sinuated radial ribs, 89
on last complete whorl, whose interstices are 2-3 times their width.
Microsculpture with 2 elements: a greatly reduced microlattice visi-
ble on a few areas under 96 x magnification and a much more promi-
nent sculpture of rather widely spaced spiral cords (fig. 850. Sutures
channeled above on postapical spire and body whorl, whorls strongly
rounded above, slightly flattened laterally above and below rounded
periphery, with evenly rounded baso-umbilical margin. Very faint
brownish yellow cast to entire shell. Umbilicus cup-shaped, widely
opened, last whorl decoiling more rapidly, contained 3.56 times in
diameter. Last '/> whorl missing, hence status of any columellar or
palatal dentition unknown. Parietal barriers 2, extending pos-
teriorly about 3 Ae of a whorl: upper broken off for entire length, only
remnants of base present; lower a broad, high, humped lamella with
irregularly gradual anterior descension and even, rather steep, pos-
terior descension. Aperture of whole specimen probably inclined
about 15 from shell axis. Height of holotype 0.86 mm., diameter 1.88
mm.
Holotype. Marshall Islands: Eniwetok Atoll, En-
gebi Islet, drill hole K-1B at 820-831 ft. depth
(Miocene #). USNM 562091.
Remarks. The spiral apical sculpture, secondary
microsculpture of strong spiral cords on the lower
whorls, and shape of the remaining parietal barriers
immediately place Vatusila eniwetokensis in the
Charopidae.
Association with recent species is handicapped by
the absence of the last half-whorl and resulting loss of
any palatal barriers. The combination of 12 apical spi-
ral ribs and a secondary microsculpture of very promi-
nent and widely spaced spiral cords is relatively
unusual in the present-day Micronesian charopid
fauna. Taxa with approximately these features include
only the Palline notera complex from Palau and the
Vatusila radiation from Fiji (Nayau, Lau Group),
Tonga (Eua), Niue, and Vaitupu, Ellice Islands. The
Palline notera complex has simple parietal barriers
(probably reduced), whereas Vatusila shows a tendency
for fusion and reduction of two parietals. The basic
shape of the parietals in V. tongensis is the same as in
V. eniwetokensis. The latter could be transformed into
the former by partial parietal fusion. Vatusila tongen-
sis, V. kondoi, and V. nayauana share the very strong
microsculpture of V. eniwetokensis. There are no fea-
tures allying it to other genera, and I have no hesita-
tion in placing it with Vatusila.
196
SOLEM: ENDODONTOID LAND SNAILS
Vatusila tongensis, new species. Figures 83a-c,
84a-b.
Diagnosis. Shell large, diameter 2.32-2.75 mm. (mean 2.53
mm.), with 4'A-5 tightly coiled whorls. Apex and spire moderately
and evenly elevated, body whorl descending much more rapidly,
spire protrusion more than Vt body whorl width, H/D ratio 0.472-
0.650 (mean 0.538). Apical sculpture of 8-12 (mean 10.4) narrow but
prominent spiral cords whose interstices are 3-4 times their width.
Postnuclear sculpture of high, narrow, rather widely spaced, protrac-
tively sinuated radial ribs, 52-72 (mean 61) on the body whorl,
whose interstices are 4-6 times their width. Ribs/mm. 6.89-8.71
(mean 7.66). Microsculpture of extremely fine radial riblets, more
than 12 between each pair of major ribs, even finer and more crowded
spiral riblets, with a secondary microsculpture of prominent spiral
cords about twice as large as the microradials, whose interstices are
1-3 times their width. Umbilicus broadly open, U-shaped, regularly
decoiling, contained 3.00-4.00 times (mean 3.31) in the diameter,
margins rounded. Sutures deep, whorls strongly rounded above,
markedly compressed laterally and slightly on basal margin. Aper-
ture subcircular. inclined about 10 from shell axis. Parietal wall
with single bifurcated barrier, extending posteriorly more than 3 Ae of
a whorl: upper blade very high and crescentic, gradual posterior
descension, rather sharp anterior descension, lower blade shorter
posteriorly, elevated medially with same anterior descension found
in Graeffedon. Both blades mounted on a broad callus that tapers
laterally beyond anterior end of bifurcated blade. Columellar wall
without barriers. Palatal barriers 2, rather deeply recessed, bulky,
extending about '/a whorl posteriorly: 1st basal in position, slightly to
strongly bifurcated, occasionally split into two parts, inner a broad,
high, crescentic, rounded, knoblike protuberance usually connected
by a callus to a much lower, cordlike blade; 2nd on outer palatal wall,
slightly supraperipheral and subperipheral to completely sub-
peripheral, about 0.25-0.36 mm. wide, crescentic in outline, with
rather gradual anterior descension, sometimes slightly concave on
upper surface with threadlike lamellae.
Vatusila tongensis is immediately recognizable by
its very large, bifurcated parietal and two very broad
palatal barriers. All other Vatusila have very slender
palatals. No other Pacific Island endodontoids match
the palatal barriers. Vatusila vaitupuensis and V.
niueana both have a less protruded spire, much more
crowded radial ribbing, and much less conspicuous
apertural barriers.
Description. Shell large, with 4 7 /s tightly coiled whorls. Apex
and spire very slightly elevated, last whorl descending moderately,
H/D ratio 0.519. Embryonic whorls l'/2, sculpture mainly eroded with
only traces of spiral cording remaining in sutures. Postnuclear
whorls with sculpture of widely spaced, high, protractively sinuated
radial ribs, rounded above, 72 on the body whorl, whose interstices
are 3-5 times their width. Microsculpture of very low and faint ra-
dial and spiral riblets crossing much more prominent, rather widely
spaced secondary spiral cords. Sutures deep, whorls slightly shoul-
dered above, laterally compressed on outer margins. Umbilicus
broadly U-shaped, regularly decoiling, contained 3.00 times in the
diameter. Color light yellow-white with broad, somewhat vague
flammulations that follow lines of growth. Aperture ovate, flattened
laterally, inclined about 5 from shell axis. Parietal wall with a
large, crescent-shaped bifid barrier, extending about '/s whorl,
structure as in diagnosis. Columellar wall without barriers. Palatal
wall with 2 oddly shaped, recessed barriers: lower a moderately high,
irregularly shaped knoblike protuberance with much lower recessed
ridgelike outer portion; outer a low, very broad, 0.36 mm. wide cal-
lus, extending about VLB of a whorl. All barriers are very minutely
serrated on all portions. Height of holotype 1.38 mm., diameter 2.67
mm.
Holotype. Tonga: Eua, hillside on cliffs east of
Ohunua, 2 miles inland at 390 ft. elevation. Collected
by H. S. Ladd on May 10, 1928. BPBM 87705.
Range. Eua Island, Tonga.
Paratypes. Eua: cliffs east of Ohunua, 2 miles in-
land at 390 ft. elevation (9 specimens, BPBM 87705,
BPBM 87660); on cliff, V 2 mile inland from Vaigana at
760 ft. elevation (9 specimens, BPBM 87632, BPBM
87693-4); main range on east side (Station T-22) at
1,000 ft. elevation (1 specimen, FMNH 152376).
Remarks. The very peculiar barriers of Vatusila
tongensis immediately identified this species. The bifid
parietal probably originated through partial fusion of
two separate barriers, a process that seems to have
been carried further in the Ellice Island V. uaitupuen-
sis and the Fijian V. kondoi. The palatal barriers vary
considerably, the lower one usually being low and
ridgelike with a lateral fanglike barrier, but often as-
suming the knoblike form of the type. The outer
palatal varies from a flattened, even callus as in the
type, to a slanted, basal callus with three or four low,
threadlike lamellae on top of it, but generally is with-
out distinct ribbing above.
Fragmentary soft parts were available and are de-
scribed below.
Description of soft parts. Several fragmentary examples were
seen, three of which had the penis and epiphallus present. Vas defer-
ens (fig. 84a, VD) reflexing at penioviducal angle, entering expanded
head of epiphallus (E). Latter more than % length of penis, entering
latter through fan of penial retractor muscle (PR). Penis 0.9-1.0 mm.
long, expanded apically, a strong median bulge, tapering gradually
to rather long and slender atrium (Y). Interior (fig. 84b) of penis with
a circularly bilobed verge and superior epiphallic pore, biscuit-like
stimulator very large and with narrow central crack.
(Based on BPBM 87660 and BPBM 87705, fragmentary parts.)
Vatusila vaitupuensis, new species. Figure 85a-d.
Diagnosis. Vatusila vaitupuensis has 1 parietal and 2 small
palatal barriers, a wider umbilicus, and much more numerous and
crowded radial ribs than the Tongan V. tongensis. Vatusila niueana
has a narrower umbilicus, only traces of apertural barriers, and
many more radial ribs than V. vaitupuensis.
Description. Shell large, with over 4'/2 relatively tightly coiled
whorls. Apex and spire barely protruding above body whorl, last
whorl descending slightly more rapidly, spire protrusion about Ve
body whorl width, H/D ratio 0.487. Apical whorls 1%, sculpture of 8
relatively broad, prominent spiral cords, whose interstices are about
2*/2 times their width. Postnuclear whorls with low, rounded,
crowded, protractively sinuated radial ribs, 115 on the body whorl,
whose interstices are 2-3 times their width. Microsculpture of fine
radial riblets, 3-5 between each pair of major ribs and very incon-
spicuous crowded spiral riblets. No secondary spiral cording. Sutures
relatively shallow, whorls slightly shouldered above, almost evenly
rounded on outer margins, very slightly compressed laterally above
and below periphery. Umbilicus broadly U-shaped, regularly decoil-
ing, contained 2.89 times in the diameter, margins rounded. Color
yellowish white with relatively regular, broad, reddish flammula-
tions that parallel lines of growth. Aperture circular, inclined about
5 from the shell axis. Parietal wall with 1 deeply recessed, low,
crescentic barrier (fig. 85d), extending less than 'Ae of a whorl. Col-
umellar wall without barriers. Palatal barriers 2: lower a short,
broad, cordlike ridge near baso-columellar margin; upper slightly
supraperipheral, moderately recessed, extending about Vs whorl, a
low, V-shaped ridge. Height of holotype 1.25 mm., diameter 2.57 mm.
Holotype. Ellice Islands: Vaitupu. Collected by L.
Isaacs in May, 1941. BPBM 189681.
Range. Vaitupu, Ellice Islands.
FIG. 83. a-c, Vatusila tongensis, new species. East of Ohunua, Eua, Tonga. Holotype. BPBM 87705; d-f, Vatusila niueana, new species.
Alofi, Niue. Holotype. BPBM 194517. Scale lines equal 1 mm. (a-c, MM; d-f, SH).
197
198
SOLEM: ENDODONTOID LAND SNAILS
PR
VD
VD
FIG. 84. Partial anatomy of Vatusila tongensis, new species. Eua,
Tonga. BPBM 87660, BPBM 87705: a, penis and epiphallus; b, vergic
papilla and stimulatory pad as seen by opening penis at junction
between collar and pad. Scale line equals 1 mm. (MM).
Paratype. Vaitupu (1 specimen, BPBM 189681).
Remarks. Only the adult holotype and a single
juvenile specimen are known. The barriers in the latter
are relatively more prominent than in the adult, but
occupy the same position and have the same shape. The
parietal (fig. 85d) closely resembles the upper blade of
V. tongensis in outline (fig. 83b), but is reduced in size.
Similarly, the palatal barriers in V. vaitupuensis oc-
cupy the same positions as the palatals of V. tongensis,
but are greatly reduced in prominence.
Vatusila niueana, new species. Figure 83d-f.
Diagnosis. Vatusila niueana has the apertural barriers re-
duced to barely visible traces. It differs from V. vaitupuensis (fig.
85a-d) also in having secondary spiral cording, a much narrower
umbilicus, and fewer, less crowded radial ribs. Vatusila tongensis
(fig. 83a-c) has very large apertural barriers, much more widely
spaced radial ribbing, and a distinctly more elevated spire.
Differences from geographically adjacent Sinployea are discussed
below.
Description. Shell large, with 4V4 rather tightly coiled whorls.
Apex and spire moderately and almost evenly elevated, last whorl
descending much more rapidly, spire protrusion more than l k body
whorl width, H/D ratio 0.500. Apical whorls l'/2, sculpture of 10 rela-
tively prominent spiral cords whose interstices are about 3 times
their width. Postnuclear whorls with prominent, rounded, rather
closely spaced, protractively sinuated radial ribs, 89 on the body
whorl, whose interstices are 2-3 times their width. Ribs/mm. 11.60.
Microsculpture of extremely fine radial riblets, 5-10 between each
pair of major ribs, slightly finer spiral riblets, and weak, rather
widely spaced secondary spiral cording. Sutures moderately im-
pressed, whorls evenly rounded on outer margins. Umbilicus open,
broadly U-shaped, last whorl decoiling slightly more rapidly, con-
tained 3.70 times in the diameter, margins rounded. Color light
yellow-white, with a few irregular, reddish flammulations that fade
out below periphery. Aperture circular, inclined about 10 from shell
axis. Parietal wall with low, broadly angled, slightly submedial
swelling, visible from normal side view only as an irregularity in
outline (fig. 83e). Columellar, basal, and lower palatal wall with a
white callus, narrower and higher at first, becoming broader, lower
and more deeply recessed up to periphery of palatal wall. Height of
holotype 1.27 mm., diameter 2.44 mm.
Holotype. Niue Island: Alofi Village at 20 m. ele-
vation. Collected on Pilea microphylla Liebm. by T. G.
Yuncker on February 12, 1940. BPBM 194517.
Range. Niue Island.
Remarks. Even with reasonably careful inspec-
tion, Vatusila niueana (fig. 83d-f) can be misidentified
as a Sinployea. The parietal swelling and palatal callus
are visible only by tilting and use of oblique lighting
into the aperture. I consider it to represent another
stage in barrier reduction within the genus Vatusila.
Although the shell measurements fall just within the
range of Sinployea uicaria and S. ellicensis (table
XXIX), in form and whorl contour V. niueana is much
closer to the other Vatusila, the Sinployea showing lat-
eral whorl flattening and a higher spire protrusion
(uicaria, fig. 58b) or much narrower umbilicus (ellicen-
sis, fig. 62c). Unfortunately, only the single example,
with no soft parts present, was available.
a be
FIG. 85. a-d, Vatusila vaitupuensis, new species. Vaitupu, Ellice Islands. Holotype. BPBM 189681: a-c, entire shell, d, parietal barrier; e-f,
Vatusila eniwetokensis (Ladd). Drillhole K-lb at 820-831 ft. depth, Engebi Islet, Eniwetok, Marshall Islands. Holotype. USNM 562091. e, lateral
view of parietal barriers;/; diagram of microsculpture. Scale line equals 1 mm.; d-f greatly enlarged. (MM).
199
200
SOLEM: ENDODONTOID LAND SNAILS
Genus Graeffedon, new genus
Shell large, with 4V&-5V4 normally coiled whorls. Apex and spire
flat to slightly elevated, body whorl very slightly to moderately de-
scending. Apical sculpture of 1 1- 13 narrow, prominent spiral cords.
Postnuclear sculpture of medium to very prominent, protractively
sinuated radial ribs, with fine radial and finer microspiral elements,
but no spiral cording. Periphery rounded, body whorl slightly flat-
tened laterally above periphery. Umbilicus open, regularly decoiling.
Color flammulations zigzagged. Parietal wall with 2 barriers, ex-
tending posteriorly about Vn whorl, upper with rather sharp anterior
descension, lower with more gradual anterior descension. Columel-
lar wall with or without a low threadlike ridge parallel to plane of
coiling, recessed or reaching lip margin. Palatal barriers 4 to 7,
short, high, almost reaching lip edge, sometimes with weak acces-
sory traces. Pallial region with rectal kidney lobe much larger and
longer than pericardia!, a distinct intrusion of mantle gland tissue
onto pallial roof. Apical genitalia apparently without unusual fea-
tures. Epiphallus with simple entrance of vas deferens, followed by 2
high pilasters. Penis with an apical muscle ring, and 3 large pilas-
ters, 1 probably acting as a vergic papilla. Lower female tract with
simple pore opening from free oviduct into common chamber of vag-
ina and spermatheca, which is lined with high, longitudinal pilas-
ters.
Type species. Endodonta graeffei Mousson, 1869.
The most similar shells are those of Semperdon,
particularly examples of S. heptaptychius (Quadras &
Mollendorff, 1894) (fig. 99a-c, e) and S. rotanus (fig.
99d). Their penial anatomy (fig. lOOb, d-g, k) is so
totally different from that of Graeffedon (fig. 87b c)
that the shell similarities clearly are convergent. Even
optical microscopic examination of the apertural bar-
riers shows that there are differences in microarma-
ture between the two genera, and the SEM photos of
Graeffedon (fig. 7a-c) and Semperdon xyleborus (fig.
8a-e) demonstrate that they are formed quite
differently (see pp. 19-20).
Graeffedon stands quite isolated from other Pacific
Island Charopinae in terms of anatomical structure.
The simple vas deferens-epiphallus juncture, very
large penial pilasters, and strong pilasters within the
vaginal-spermathecal chamber are quite different from
those found in the typical Charopinae. It is quite possi-
ble that eventual subfamily separation of Graeffedon
will be required, but until many more extralimital
taxa have been studied, retention of Graeffedon within
a broadly defined Charopinae is preferred.
The only genus with which I can at present relate
Graeffedon in terms of anatomy is Helenoconcha
Pilsbry, 1892, from St. Helena in the South Atlantic.
The only dissected species, Helenoconcha relicta Solem
(1977c, figs. 184a-b, 185a-d), agrees in pallial
structure (except for lacking the mantle gland intru-
sion) and has exactly the same type of penial structure.
The two taxa differ only in details of pilaster position
and size, but the penis complex of Helenoconcha is re-
markably similar to Graeffedon. At present the inter-
nal structures of the terminal genitalia in the South
African endodontoid taxa, Trachycystis Pilsbry, 1893,
and Afrodonta Melville & Ponsonby, 1908, are un-
known (see Solem, 1970c), but the external genital fea-
tures of Trachycystis illustrated by Watson (1934, pis.
20-21) are compatible with at least the latter genus
being a member of this complex.
It is thus premature to attempt formal designation
of a subfamily unit for Graeffedon.
Specimens of Graeffedon are rare in collections
and in the field. The three species differ in size,
sculpture, and details of the palatal barriers (see Diag-
noses). Because of their allopatric distribution, no
structural key is presented. Their known distributions
are:
Graeffedon graeffei Upolu, Samoa
G. sauaiiensis Savai'i, Samoa
G. pricei Tongatapu, Tonga.
Graeffedon graeffei (Mousson, 1869). Figures 7a-c,
86a-c, 87a-d.
Endodonta graeffei Mousson, 1869, J. de Conchyl. 17, pp. 332-333,
pi. 14, fig. 3 Upolu, Samoa.
Pitys graeffei (Mousson), Pease, 1872, Proc. Zool. Soc. London,
1871, p. 474; Garrett, 1887, Proc. Acad. Nat. Sci. Philadelphia,
1887, pp. 130-131.
Helix (Endodonta) graeffei (Mousson), Pfeiffer, 1876, Monog. helic.
viv., 7, p. 258; Tryon, 1887, Man. Conchol., (2) 3, p. 65, pi. 12, fig.
31.
Endodonta (Thaumatodon) graeffei (Mousson), Pilsbry, 1893, Man.
Conchol., (2)9, p. 27.
Diagnosis. Shell very large, diameter 4.61-5.92 mm. (mean
5.23 mm.), with 4%-5'/4 rather tightly coiled whorls. Apex and early
spire flat or barely protruding, lower spire descending slightly, body
whorl much more rapidly, spire protrusion about J Ai body whorl
width, H/D ratio 0.472-0.560 (mean 0.525). Apical sculpture of
11-13 (mean 12.0) narrow but prominent spiral cords whose in-
terstices are 4-5 times their width. Postnuclear sculpture of high,
prominent, V-shaped to U-shaped, crowded, slightly protractively
sinuated radial ribs, 92-121 (mean 105.0) on the body whorl, whose
interstices are 2-3 times their width. Ribs/mm. 6.27-6.51 (mean
6.39). Microsculpture of proportionately fine radial riblets, 5-8 be-
tween each pair of major ribs, crossed by much finer and more
crowded spiral riblets. No secondary spiral cording. Umbilicus mod-
erately open, V-shaped, regularly decoiling, contained 3.53-4.19
times (mean 3.94) in the diameter, margins rounded. Whorls dis-
tinctly flattened laterally above periphery and on basal margin,
aperture inclined about 15 from shell axis. Parietal barriers 2, ex-
tending posteriorly slightly more than VH whorl: upper very high and
broadly expanded above posteriorly, this portion deflected down-
ward, then gradually descending and narrowing to anterior quarter,
then abruptly descending on anterior 8th; lower broadly expanded,
serrated, and crescentic on posterior half, narrowing above and with
even descension anteriorly. Columellar wall with (40% ) or without
(60% ) a low, broadly rounded, high, threadlike ridge, lying parallel
to plane of coiling and reaching lip margin. Palatal barriers 5, high
and prominent, reaching almost to lip margin: lower 4 progressively
higher, longer, more slender, changing from abrupt to more gradual
anterior descension, all crescentic, with height about % length, ex-
tending posteriorly less than Vs whorl, with 1st at basal-columellar
margin; 4th subperipheral; lower 2 have anterior edge turned inward
just before reaching lip; upper supraperipheral, greatly reduced in
height, a bladelike barrier with equally gradual anterior and pos-
terior descension, longer than 4th, and slightly recessed within aper-
ture.
Graeffedon graeffei is much larger than the Savai'i
Island G. sauaiiensis or the Tongan G. pricei (table
XLIII). The former has a much wider umbilicus, seven
palatal barriers, finer sculpture, an elevated apex, and
more rounded whorls; the latter has fewer and more
prominent radial ribs, only four palatal barriers, a flat
apex, and is smaller in size.
SYSTEMATIC REVIEW
201
a
FIG. 86. a-c, Graeffedon graeffei (Mousson). Upolu, Samoa. Lec-
totype. Zoologisches Institut der Universitat Zurich. Scale line
equals 1 mm. (SG).
Description. Shell very large, with slightly more than 4% mod-
erately tightly coiled whorls. Apex and early spire flat, later whorls
descending a little, body whorl more rapidly, H/D ratio 0.527. Apical
whorls 1%, sculpture of 12 narrow cords. Postnuclear sculpture of
high, prominent, crowded, protractively sinuated radial ribs, 98 on
the body whorl, whose interstices are 2-3 times their width. Micro-
sculpture a lattice of fine, relatively crowded and numerous radial
riblets, 5-8 between each pair of major ribs, crossed by much finer
and more crowded spiral riblets. Sutures deeply impressed, whorls
strongly rounded above, somewhat flattened laterally above periph-
ery and slightly on basal margin. Umbilicus moderately open, V-
shaped, regularly decoiling, contained 3.93 times in the diameter.
Color light yellowish white with very prominent zigzag reddish
brown flammulations. Aperture subovate, flattened laterally above
periphery, inclined about 15 from the shell axis. Parietal barriers 2,
extending a little more than '/ whorl: upper high, bladelike, slightly
expanded above posteriorly with relatively sharp anterior descen-
sion; lower more crescentic, more broadly expanded above with com-
paratively gradual anterior descension. Columellar wall without
barriers. Palatal barriers 5, reaching lip margin, short and crescen-
tic: lower at baso-columellar margin, sharply descending to lip edge;
2nd, 3rd, and 4th crescentic, very high, slightly expanded above, 2nd
reaching lip margin with very sharp anterior descension, 3rd slightly
recessed with more gradual descension, 4th a little more recessed
with much more gradual anterior descension; upper a longer, lower,
ridgelike lamella, narrow above with very gradual posterior rise and
anterior descension. All barriers minutely pustulose or serrated
above. Basal and columellar portion of lip with a low, broad, white
callus. Height of holotype 2.60 mm., diameter 4.94 mm.
Holotype. Samoa: Upolu. Collected by E. GraefFe.
Zoologisches Museum der Universitat, Zurich.
Range. Probably the uplands of Upolu, Western
Samoa.
Material. Upolu (4 specimens, Zurich, Brussels,
FMNH 116976, FMNH 147544): summit of Mt. Solaua
(Station 39) under ground debris in heavy forest at
2,000 ft. elevation (1 specimen, FMNH 153420).
Remarks. Besides the type in the Mousson col-
lection, two specimens were found in museums iden-
tified as Sinployea complementaria (Mousson, 1865). A
single example was collected under rotting wood in
1965 together with 18 specimens of S. complementaria.
In size and general appearance (figs. 56a-f, 86a-c) the
two species are very similar, with the elevated spire
and narrower umbilicus of S. complementaria (fig.
56a-f) not being obvious during field examination. Ex-
ternally the animal of Graeffedon graeffei differs in
having the body without any grayish markings on the
neck and sides of the foot. The absence of apertural
barriers in S. complementaria and their presence in G.
graeffei immediately separate the two species.
In order to dissect the one specimen, partial de-
struction of the shell was necessary. The soft parts
broke during extraction, but reasonably full study of
the anatomy was possible. The significant differences
have been discussed previously (p. 29).
Description of soft parts. Foot and tail completely retracted
into shell. Sole and pedal grooves typical, no caudal horn or middor-
sal groove developed. Slime network conspicuous. Body color
yellow-white, no trace of darker markings.
Mantle collar (MC) partly shredded during extraction because of
large barriers, a prominent glandular extension (MG) onto pallial
roof. Anus (A) opening slightly in front of external ureteric pore
(KX), vague grooves leading through pneumostomal opening.
202
SOLEM: ENDODONTOID LAND SNAILS
TABLE XLIII. - RANGE OF VARIATION IN GRAEFFEDON.
NUMBER OF
SPECIMENS
graeffei
(Mousson)
savailensis
price!
RIBS
105(92-121)
88
68
RIBS/MM.
6.39(6.27-6.51)
H/D RATIO
graef. 0.525(0.472-0.560)
savai. 0.485
pricei 0.485
HEIGHT
2.75(2.37-3.26)
1.61
1.61
DIAMETER
5.23(4.61-5.92)
3.32
3.32
APICAL
WHORLS UMBILICUS D/U RATIO CORDS
5-(4 3/4-5 1/4) 1.36(1.15-1.48) 3.94(3.53-4.19) 12.0(11-13)
4 1/4 1.02 3.26 11
4 1/8- 0.92 3.61 WORN
SPIRE
ELEVATION
graef. 0.17(0.10-0.20)
saval. 0.15
pricei FLAT
BODY WHORL
WIDTH
1.83(1.68-2.04)
1.05
1.13
SP/BWW PR
0.092(0.058-0.118) 2
0.141 2
2
P
5
7
4+2
Pallial region (fig. 87a) extending about 5.5 mm., distance from
anus to anterior tip of kidney 1.65 mm. Lung roof clear, without
granulations. Kidney (K) bilobed, pericardia! lobe less than half
length of rectal lobe, very narrow, rectal lobe crossing hindgut into
parietal wall, length 3.85 mm. Ureter (KD) with short primary arm
jammed between lobes of kidney with no lung roof visible, secondary
ureter widening along hindgut. Heart (H) angled to hindgut, slightly
more than half length of pericardia! kidney lobe. Principal pulmo-
nary vein (HV) unbranched and inconspicuous, reaching to edge of
mantle collar extension. Hindgut (HG) extending well past pallial
cavity apex before leaving parietal-palatal margin.
Ovotestis (fig. 87d, G) a single clump of large, long acini, pal-
mately clavate, lying parallel to whorl sides above stomach apex.
Hermaphroditic duct (GD) long, expanded evenly for most of length,
abruptly narrowing before reflexing to enter base of globular talon.
Albumen gland completely destroyed by parasites in dissected
example. Talon (not shown) large, globular, with very narrow neck to
carrefour. Prostate (fig. 87b, DG) of large acini opening into groove in
inner uterine wall, variously folded. Uterus (UT) bipartite, upper
chamber thin-walled and narrow, lower grossly expanded and
thick-walled.
Vas deferens (VD) tapering and with glandular walls to
penioviducal angle, narrow and muscular after passing through
muscle threads, entering epiphallus without special structures.
Epiphallus (E) larger than penis, curved and twisted, internally (fig.
87c) with one very high and rounded and a second lower and hemi-
spherical pilaster that taper into epiphallic pore. Penial retractor
(PR) a broad muscle band arising from diaphragm and inserting in a
narrow line onto head of penis along middle zone of apical penial
pilaster. Penis (P) about 1.48 mm. long, sharply constricted subapi-
cally, then tapering. Internally (fig. 87c) with a dome-shaped pilaster
and epiphallic pore (EP) on one side of center occupying area above
constriction, below with a huge spade-shaped pilaster tapering to
atrium. Lateral to this a "hoe"-shaped pilaster tapering to a broad
glandular ridge and a high, longitudinal pilaster tapering into glan-
dular area. Atrium (Y) short, without marked features.
Free oviduct (UV) muscular, short, with glandular walls. Sper-
matheca (S) with expanded head next to albumen gland, slender
shaft bound to prostate-uterus, expanded basal portion with 2 large
and several smaller tapered pilasters (fig. 87c). Vagina (V) with
many narrow pilasters, externally with numerous muscle strands
holding it to the body wall.
Esophagus distended, typical in position, stomach occupying %
of a whorl. Intestinal looping without unusual features. Digestive
glands mostly replaced by parasites above stomach apex, partly in
region of esophagus-stomach junction. Salivary glands united above
esophagus, right gland greatly reduced in size.
(Based on 1 adult, FMNH 153420.)
Graeffedon savaiiensis, new species. Figure 88a-c.
Diagnosis. Graeffedon sauaiiensis differs from the Upolu G.
graeffei (Mousson, 1869) in its slightly and regularly elevated apex,
possession of 7 palatal barriers, wider umbilicus, and much smaller
size. The Tongan G. pricei (fig. 88d-f) has a flat apex and spire, much
coarser radial ribbing, lower and shorter parietals, and only 4 major
palatals, although essentially identical in size and proportions.
Description. Shell slightly larger than average, with 4V* nor-
mally coiled whorls. Apex and spire slightly and evenly elevated, last
whorl not descending more rapidly, spire protrusion about l h body
whorl width, H/D ratio 0.485. Apical whorls 1V4, sculpture of 11 thin,
relatively high, spiral cords. Postnuclear whorls with thin, lamel-
late, protractively sinuated radial ribs, 88 on the body whorl, whose
interstices are 2-4 times their width. Ribs/mm. 8.43. Microsculpture
of fine radial riblets, 3-6 between each pair of major ribs, with much
finer and more crowded spiral riblets. Sutures moderately impressed,
whorls strongly rounded above suture, slightly flattened laterally
and on basal margin with evenly rounded periphery. Umbilicus
widely open, V-shaped, last whorl decoiling more rapidly, contained
3.26 times in the diameter. Color light yellow-white with irregular
reddish brown flammulations becoming darker on body whorl. Aper-
ture ovate, slightly flattened laterally above periphery, inclined
about 10 from shell axis. Parietal barriers 2, extending posteriorly
% whorl: upper very high and crescentic posteriorly, serrated and
expanded above, flattening out anteriorly, then descending rather
sharply on anterior 8th; 2nd a much lower, bladelike lamella,
broadly expanded above and slightly elevated posteriorly, with
gradual descension over anterior half. Columellar wall with single,
H
VD
UV
FIG. 87. Anatomy of the Samoan Graeffedon graeffei (Mousson). Station 39, Mt. Solaua, Upolu, Samoa. FMNH 153420: a, pallial region; b,
genitalia; c, interior of penis, epiphallus, spermathecal base; d, ovotestis and hermaphroditic duct. Scale lines equal 1 mm. (SH).
203
def
FIG. 88. a-c, Graeffedon savaiiensis, new species. Mataulano Lake, Savai'i, Samoa. Holotype. BPBM 95738; d-f, Graeffedon pricei, new
species. Manima Id., Tongatapu, Tonga. Holotype. FMNH 152399. Scale lines equal 1 mm. (a-c, MM; d-f, SH).
204
SYSTEMATIC REVIEW
205
low, threadlike ridge, lying parallel to plane of coiling, reaching top
of columellar callus. Palatal barriers 7, short, high, reaching lip
margin for early ones, then later barriers slightly recessed: lower at
baso-columellar margin, hemispherical, very high, with abrupt an-
terior descension, curving back inward at lower anterior margin;
3rd, 4th, and 5th progressively lower, longer, less expanded above,
changing to more gradual anterior descension with 5th crescentic, all
subperipheral; 6th equal in height to 5th, slightly supraperipheral,
with more gradual anterior descension; 7th high, bladelike, much
lower than 6th, slightly concave above, moderately expanded, with
gradual anterior descension, injured anteriorly, slightly recessed
within aperture. All palatals weakly serrated on upper edge and
sides. Height of holotype 1.61 mm., diameter 3.32 mm.
Holotype. Samoa: Savai'i, near Mataulano Lake.
Collected in leaf mould on a hillside by E. Christopher-
son on October 2-3, 1929. BPBM 95758.
Range. Interior of Savai'i, Western Samoa.
Material. Known only from the holotype.
Remarks. The elevation of Mt. Mataulano is
2,952 ft., but I have no data as to the elevation of the
lake at which this species was collected. The single
shell is probably subadult, but the distinctness from G.
graeffei is based on the former having smaller nuclear
whorls, a flat apex and early spire, wider and higher
major radial ribs, and a noticeably narrower umbilicus.
Shape of the parietal barriers is identical, but in G.
savaiiensis they are shorter and proportionately
higher. Besides the increased number of palatals, in G.
savaiiensis they are shorter and hemispherical in out-
line, rather than being crescentic. The columellar in G.
savaiiensis is lower and more deeply recessed.
Despite particular effort by Mr. Price, no speci-
mens were collected on Savai'i in 1965.
Graeffedon pricei, new species. Figure 88d-f.
Diagnosis. Graeffedon pricei differs from the Savai'i G.
savaiiensis in its flat apex and spire, heavier and less crowded rib-
bing, smaller parietals, and presence of only 4 palatals. Graeffedon
graeffei from Upolu is much larger, has the body whorl descending
much more rapidly, and has proportionately much larger apertural
barriers.
Description. Shell slightly larger than average, with a little
less than 4'/s rather tightly coiled whorls. Apex and spire flat, body
whorl descending slightly, H/D ratio 0.485. Apical whorls 1%,
sculpture eroded except for traces of narrow spiral cords in sutures
and umbilicus. Postnuclear whorls with very prominent, wide, pro-
tractively sinuated radial ribs, 68 on the body whorl, whose in-
terstices usually are 2-4 times their width. Microsculpture mostly
obscured, visible on spots as consisting of very fine radial riblets,
6-12 between each pair of major ribs, crossed by slightly finer and
more crowded spiral riblets. Sutures impressed, whorls strongly
rounded above and on periphery, slightly compressed laterally above
and below periphery. Color leached from shell, with faint traces of
reddish flammulations visible above periphery. Umbilicus open,
broadly V-shaped, regularly decoiling, contained 3.61 times in the
diameter. Aperture circular, slightly compressed laterally above and
below rounded periphery, inclined about 10 from shell axis. Parietal
barriers 2, relatively low, extending posteriorly '/* whorl: upper ele-
vated on posterior quarter, expanded and serrated above, gradually
descending to anterior threadlike quarter; 2nd equal in height,
slightly more expanded above on elevated posterior quarter, rather
sharply descending to midpoint, anterior half a high threadlike
ridge. Columellar wall without barriers. Palatal barriers 4, high,
bulbously expanded above, short, reaching lip edge, with two thread-
like accessory traces: lower at baso-columellar margin, hemispheri-
cal, anterior descension abrupt, margin deflected inward just before
lip; 2nd slightly higher, equally expanded above, with sharp anterior
descension, but no inward deflection; 3rd equal in height, slightly
subperipheral, with more gradual anterior descension; 4th supra-
peripheral, reduced in height, a bladelike lamella strongly expanded
above, slightly recessed, with gradual anterior descension. Accessory
traces low and threadlike, located midway between 1st and 2nd, 2nd
and 3rd palatals, slightly recessed, shorter than major palatals.
Height of holotype 1.61 mm., diameter 3.32 mm.
Holotype. Tonga: Tongatapu, Manima Island, 4
miles northeast (Station T-4) of Nuku'alofa in second
growth on a coral atoll. Collected by Laurie Price on
January 26, 1966. FMNH 152399.
Range. Probably Tongatapu and its satellite is-
lands, Tonga.
Material. Only the holotype is known.
Remarks. Graeffedon pricei is identical in height
and diameter to the single known example of G.
savaiiensis, but readily separable by the characters
cited in the diagnosis. Its aperture is more evenly
rounded, and the ribbing difference is particularly ob-
vious. The position of the major palatals and two acces-
sory palatal traces in G. pricei does not correspond to
the position of the palatals in G. savaiiensis or G.
graeffei.
Collections on Tongatapu were made during a se-
vere drought. No living endodontids were collected and
comparatively little live material in other groups.
Great pleasure is taken in naming this species after
Mr. Laurie Price of Kaitaia, New Zealand, in grateful
recognition of his collecting activities for Field
Museum of Natural History.
Subfamily TRUKCHAROPINAE, new subfamily
Formal description of this subfamily has been
given on p. 70.
Several genera confined to the Palau and Caroline
Islands show major anatomical modifications. In sharp
contrast to the Charopinae and Semperdoninae, there
is no indication of differentiation into a vas deferens
and an epiphallus. The vas starts as a thin-walled
transparent tube that rapidly changes into the typi-
cally muscular shiny state seen in almost all endodon-
toid species. There may be slight diameter changes at
the penioviducal angle, but usually the tube retains a
constant diameter until entering the penis. Except in
Jokajdon, these genera all agree in having the vas def-
erens tranverse the penis head, then passing through
the penial retractor muscle fan before entering the
penis. In both the Charopinae and Semperdoninae,
there is a prominent epiphallus developed, with a com-
plicated valvular entrance present in many
Charopinae (figs. 29i-j, 57e).
Although the Palau and Caroline Island genera
grouped as the Trukcharopinae share this basic pat-
tern of vas deferens-penis junction, there is much
greater diversity in internal penial structure and mod-
erate divergence in other genital features. The degree
of genital difference between these genera is much
greater than that which is seen in the Pacific Island
206
SOLEM: ENDODONTOID LAND SNAILS
Charopinae and Semperdoninae, and involves both
addition and elimination of structures. Shell structure
is diverse, with an array of large and small, tightly and
loosely coiled, heavily sculptured and with reduced
sculpture, with and without barriers, elevated spires to
deeply depressed apices all represented. Pallial organs
show variation, but not to an extreme extent. Only 11
species are involved, but their range of diversity ex-
ceeds that in the other subfamilies.
At this time I prefer not to speculate on the mutual
affinities of these genera. Except for Jokajdon and
Palikirus, which could be monophyletic, the differences
are large enough that I am somewhat dubious of local
derivation. Until enough Australian and New Zealand
charopids have been dissected to establish the pattern
of variation within that complex, the possibility that
the Micronesian genera represent relicts from an early
faunal migration from Southeast Asia through New
Guinea to Australia, New Zealand, and New Caledonia
cannot be excluded. Such an early migration would be
prior to the spread of the helicarionid taxa presently
dominant in the Indonesian-New Guinea Archipelago.
Although the Semperdoninae and generalized Charo-
pinae appear monophyletic with reasonably clear pat-
terns of change, the Trukcharopinae share the lack of
an epiphallus and enfolding of the vas deferens within
the penial retractor muscle but represent a great vari-
ety of other experiments.
Major items of anatomical differences are sum-
marized in the following paragraphs. Roimontis to-
lotomensis, Palline biakensis, and Palikirus ponapicus
have not been dissected and thus are excluded from the
following discussion.
Variations in the pallial cavity involve length, de-
gree of kidney overlap onto the parietal wall, relative
length of the pericardial and rectal kidney lobes, and
shape of the kidney. Pallial cavity length is about
three-quarters of a whorl in Jokajdon (fig. 93a); one-
half whorl in Palikirus, Palline, Trukcharopa, and
Kubaryiellus; and one-quarter whorl in Russatus (fig.
90a). The square shape and equal kidney lobe length in
Russatus is correlated with the whorl count reduction
and coiling change. The other genera have elongated
kidneys with the lobes of unequal length. In Jokajdon
(fig. 93a), Palline (fig. 93e, g) and Kubaryiellus (fig.
90e) the rectal lobe is twice the length of the pericar-
dial; in Palikirus and Trukcharopa the rectal lobe is
207c-3(y7c longer than the pericardial. There is not a
direct correlation between lobe length and parietal
wall overlap, because there is moderate overlap in
Palline and Palikirus, strong overlap in Jokajdon,
slight overlap in Trukcharopa, and no overlap in
Kubaryiellus and Russatus. In Jokajdon and Palline
the pericardial kidney arm is greatly reduced in width
as well as length, but in Palikirus, with moderate kid-
ney overlap, the pericardial arm is not reduced in size.
I could not correlate the degree of overlap or differences
in lobe length with shell features such as coiling pat-
tern or body whorl width. Possibly the size factor is
critical because the three small genera, Jokajdon, Pal-
line, and Palikirus, have marked kidney overlap onto
the parietal wall, whereas the larger genera do not.
More data are needed.
Previous extraction of the soft parts in many
species meant that the apical genitalia could not be
studied. Often the apical part of the visceral hump
breaks off when the animal is pulled out. It remains in
the shell and subsequently dries up. Thus only partial
data are available on a dichotomy into species having
one or two alveolar clumps in the ovotestis. No size
correlation exists, since the very small Jokajdon and
quite \argeKubaryiellus have a single clump, whereas
the small Palline and medium-sized Trukcharopa have
two distinct clumps, one lying completely apicad of the
other, with both touching the palatal surface of the
whorl. I could not examine the ovotestis of Palikirus or
Russatus. In Jokajdon the ovotestis alveoli were fewer
in number and proportionately much larger and longer
than in the other genera. Sufficiently accurate mea-
surements for comparison of midvisceral genitalia
proportions were not obtained. I saw no qualitative or
obvious proportional differences in this area.
Distinctions are present in proportions and rela-
tive development of the free oviduct, spermatheca, and
vagina. Normally the basal portion of the sper-
mathecal shaft is greatly expanded in the Charopidae,
with the interior having various complicated glandular
or muscular pilasters and ridges (fig. 87c). Jokajdon is
radically different in lacking any expansion of the
lower spermathecal shaft (fig. 93b). It also is the only
Micronesian genus in which the vagina is distinctly
longer than the free oviduct. Russatus (fig. 90c) has a
short vagina and a short free oviduct with the base of
the spermathecal shaft globosely and suddenly ex-
panded. These features correlate with the reduced
whorl count and shortened pallial cavity, because a
zonal compaction of the midvisceral genitalia typically
accompanies such an alteration in coiling pattern. The
other four genera have rather long free oviducts. In
Palikirus the vagina is quite short; in Kubaryiellus
(fig. 90f), Trukcharopa (fig. 90i), and Palline (fig. 93f,
h) the vagina is slightly shorter than the free oviduct.
Spermathecal shaft expansion is essentially the same
in Palikirus and Kubaryiellus, with the gradual ta-
pered expansion occurring over a distance equal to the
free oviduct length. In Trukcharopa there is a rela-
tively sharp expansion that occurs over little more
than half of the free oviduct length, whereas in Palline
there is very gradual expansion that starts about at the
mid-uterine level, well above the free oviduct origin.
The functional significance of these changes is un-
known.
Fundamental differences exist in the penial com-
plex. Most of these are concerned with the functioning
surfaces of the penis, but even just external shape and
proportions show major alterations. Palline (fig. 93f, h),
Trukcharopa (fig. 90i), and Palikirus agree in having
an ovately to globosely expanded penial head following
SYSTEMATIC REVIEW
207
a much shorter, narrow tube (almost equal to head
length in Truckcharopa). Russatus is basically the
same, except that the expanded head is clearly triangu-
lar in shape (fig. 90b). Kubaryiellus (fig. 90f) has a very
long, club-shaped penis with the shaft tapering gradu-
ally to the atrium. Jokajdon (fig. 93b) has the penis
tapered apically and basally from a median expansion.
Penial size is best expressed in relation to shell diame-
ter because Russatus is 214 times the diameter of
Jokajdon. Three genera, Russatus, Palikirus, and Jo-
kajdon, have a long penial retractor muscle, and the
ratio of total penis length to shell diameter is between
0.15 and 0.30. Palline and Kubaryiellus have short pe-
nial retractor muscles, and the ratio is between 0.45
and 0.60. Trukcharopa has a short penial retractor and
total penis length almost equals the shell diameter.
Because origin of the penial retractor muscle normally
lies near the apex of the pallial cavity, increasing rela-
tive penis length shortens the penial retractor. Unlike
many Endodontidae, no Pacific Island Charopidae
have the penial retractor muscle detached from the
diaphragm and joined to the free retractor system.
Internally, Palline has a tubular verge that api-
cally may function as an epiphallic chamber; Kubar-
yiellus, a trilobed vergic papilla (fig. 90h); Russatus
(fig. 90d), a simple pore with a lateral, rather long
caecum on the inner wall of the penis head; Jokajdon,
Palikirus, and Trukcharopa (fig. 90j), a simple pore
with no major apical structures in the penis. Some form
of muscular "pocket" pilaster is found in Jokajdon,
Palikirus, and Trukcharopa, with the pilaster globular
in Palikirus, of the Sinployea-type in Jokajdon, and
greatly changed in Trukcharopa. Weak apical lon-
gitudinal pilasters are found in Jokajdon and
Palikirus, strong submedial longitudinal pilasters in
Kubaryiellus. Russatus has weak, circular glandular
pilasters, whereas Palline has a lateral pocket with
concentric glandular pilasters and a large, median
thick circular or partly circular pilaster. Jokajdon is
unique in having a separate free retractor muscle at-
tached to the penioviducal angle.
Of the above changes, the verge and circular pilas-
ters in Palline, internal caecum in Russatus,
penioviducal angle muscle in Jokajdon, and major lon-
gitudinal pilasters in Kubaryiellus represent entirely
new structures. Opening of the vas deferens into the
penis through a simple pore in Russatus, Palikirus,
Jokajdon, and Trukcharopa represents a loss of
structure when compared with the Charopinae. Other
structures could be relatively simple modifications
from the Sinployea pattern, but the items detailed
above represent loss or gain of entire structures.
Factors of shell variation are discussed in greater
detail elsewhere (pp. 8-23). Here it is necessary to
note only a few points. It is the smallest shells, Jokaj-
don and Palline, that have the tightest coiling pattern
(figs. 92a, d; 94a) and greatest development of aper-
tural barriers (fig. 92b, e). The largest shells,
Kubaryiellus and Russatus, have the loosest form of
coiling (fig. 89a-f) and totally lack apertural barriers.
Only Palline and Palikirus show the normal charopid
shell coloration of simple yellow-horn or with traces of
darker flammulations. The remaining genera have the
shell reddish (Jokajdon, Trukcharopa), reddish brown
(Kubaryiellus, Roimontis), or deep reddish purple-
brown (Russatus). Only Palline (fig. 94b) has a moder-
ately elevated spire, most of the others have it nor-
mally flat or slightly elevated, whereas in Kubaryiellus
(fig. 89b) the apex is distinctly depressed, and in
Roimontis (fig. 91a-b) it is very depressed. Umbilical
width shows only moderate variation, with slight nar-
rowing in Palline (fig. 94c) and great narrowing in
Russatus (fig. 89f). Sculpture is greatly reduced in
Russatus, greatly enlarged in Palikirus (fig. 91d-f). In-
terpretation of these variations is rendered difficult by
the high concentration of species on Ponape. Palline
notera is found on Koror, Peleliu, and Babelthuap in
the Palau Islands; Palline biakensis, on Biak Island,
West Irian; and Trukcharopa is restricted to the Truk
Group and Lukunor in the Central Carolines. The re-
maining eight species are restricted to Ponape. We
have no exact locality for Palikirus ponapicus (Mollen-
dorff, 1900), but the other seven species were taken at
one or more stations by the B. P. Bishop Museum Mic-
ronesian Expedition in early 1936. Several species live
together, because three to five species were collected at
four stations (table XLIV). No data exist concerning
any ecological separation of the species. The genital
differences, except between the two species of Jokaj-
don, cannot be ascribed to a simple character dis-
placement phenomenon. The size of the penis itself is
so different between sympatric forms, and both dele-
tion and addition of structures are involved.
Ponape offers considerable opportunities for inves-
tigation of isolating mechanisms in land snails because
the multitude of sharply differentiated charopids there
is equivalent to the remarkable radiation of trocho-
morphid genera and species, Kondoa, Hogolua, and
Brazieria in the Truk Group (Baker, 1941, pp. 272-
280), and Trochomorpha itself on Ponape (Baker, 1941,
pp. 290-296). Rapa shows an even greater radiation of
Endodontidae (Solem, 1976b, pp. 490-492), but these
taxa show far less dramatic changes in genital
structures, a much greater amount of local geographic
speciation, and probably are extinct.
Genus Trukcharopa, new genus
Shell of average size, with about 4% normally coiled whorls.
Apex and spire flat to slightly elevated, body whorl descending
slightly. Apical sculpture of about 10 prominent spiral cords. Post-
nuclear sculpture narrow, crowded, protractively sinuated radial
ribs, averaging about 18/mm. on body whorl. Microsculpture of fine
radial riblets, finer spiral, and prominent secondary spiral cords.
Umbilicus widely open, margins rounded. Whorls only slightly com-
pressed laterally above and below rounded periphery, aperture in-
clined about 20" from shell axis. No apertural barriers developed.
Pallial region extending Vi whorl apically, with rectal kidney arm
20%-30V< longer than pericardia!. Ovotestis with 2 clumps of alveoli,
1 above other, hermaphroditic duct partly convoluted. Penial retrac-
tor muscle relatively short, enfolding vas deferens before inserting
208
SOLEM: ENDODONTOID LAND SNAILS
TABLE XLIV. - SYMPATHY OF CHAROPIDAE
ON PONAPE, CAROLINE ISLANDS.
Station 91
Palikirus cosmetus
Kubaryiellus kubaryi
Jokajdon tumidulus
Jokajdon calllzonus
Station 92
Roimontis tolotomensis
Russatus nigrescens
Kubaryiellus kubaryi
Jokaj don tumidulus
Jokajdon callizonus
Station 93
Roimontis tolotomensis
Russatus nigrescens
Jokajdon tumidulus
Jokajdon callizonus
Station 139
Palikirus cosmetus
Kubaryiellus kubaryi
Jokaj don tumidulus
on penis head. Penis long, head greatly expanded, internally with a
huge medial stimulator, no vergic papilla, weak papillose glandular
walls in lower tube. Spermatheca with basal part of shaft greatly
expanded, vagina relatively long.
Type species. Trukcharopa trukana, new species.
Both Kubaryiellus and Trukcharopa have much
longer penes (fig. 90f, i) than any of the other Microne-
sian genera that lack an epiphallus, but the structures
of the penes are very different. Trukcharopa has the
head grossly expanded, lacks a vergic papilla, and has
a huge median muscular stimulatory pad on one wall
of the expanded chamber (fig. 90i-j). Serial sections of
the penial region will be required before the exact
structure of this stimulatory pad can be determined,
but the essential differences from the other Microne-
sian genera are obvious.
Trukcharopa is nearest in shell structure to the
Polynesian-Melanesian genus Sinployea. Anatomi-
cally the two differ in Sinployea having a prominent
epiphallus that is about half the penis in length, the
penis has a distinct vergic papilla, and the typical
stimulatory pad occupies the median part of the penis.
Sinployea kusaieana is the only named Micronesian
species in that genus. Conchological comparisons with
Trukcharopa trukana (tables XXVII, XLV) show that
they are virtually identical in mean diameter, umbili-
cal width, D/U ratio, and body whorl width. Sinployea
kusaieana (fig. 63a-c) is a light reddish yellow and has
an average of 91 major radial ribs, 10.3-13.1 ribs/mm.
on the body whorl, with five to 10 microradial riblets
between each pair of major ribs. In contrast, Truk-
charopa trukana (fig. 63d-f) is dark reddish brown in
color and has an average of 150 major radial ribs,
14.1-20.8 ribs/mm, on the body whorl, with only two to
four microradials between each pair of major ribs. I
could find no conchological features in Trukcharopa
that would differentiate it generically from Sinployea.
Both pallial region and male genital structures (fig.
90i-j) place T. trukana as being more closely related to
the Micronesian genera than to Sinployea.
In choosing Trukcharopa as a name for this genus,
I am reflecting both its great similarity to the tradi-
tionally named "Charopa" (=Sinployea) of Polynesia
and its geographic location.
Trukcharopa trukana, new species. Figures 63d-f,
Diagnosis. Shell smaller than average, diameter 2.50-3.08
mm. (mean 2.66 mm.), with 4'/s-4% rather tightly coiled whorls.
Apex and early spire normally flat, sometimes slightly protruding,
rarely moderately elevated, body whorl usually descending slightly
more rapidly, spire protrusion generally much less than 'Ac body
whorl width, occasionally moderately elevated, H/D ratio 0.371-
0.513 (mean 0.450). Apical sculpture of 8-12 (mean 9.68) prominent
spiral cords, whose interstices are 2-3 times their width. Postnuclear
sculpture of low, narrow, crowded, slightly protractively sinuated
radial ribs, 112-197 (mean 150.1) on the body whorl, whose in-
terstices usually are l'/2-2 times their width. Ribs/mm. 14.1-20.8
(mean 17.7). Microsculpture of very fine, rather widely spaced radial
riblets, 2-4 between each pair of major ribs, much finer and more
crowded spiral riblets, with a secondary sculpture of spiral cords that
are more prominent than the microradial riblets and whose in-
terstices are about 3-4 times their width. Umbilicus broadly open, V-
to U-shaped, regularly decoiling, contained 3.07-4.00 times (mean
3.43) in the diameter, margins evenly rounded. Sutures deep, whorls
strongly rounded above, slightly compressed laterally above
periphery and on basal margin, with evenly rounded outer and col-
umellar margins. Aperture subovate, compressed laterally above
periphery and on basal margin, inclined about 20 from shell axis. No
apertural barriers, but a weak to moderate callus extending from
columellar region up to periphery in some individuals.
Trukcharopa trukana is much smaller, has much
more crowded radial ribbing, more apical cords, and
lacks the depressed spire of Kubaryiellus kubaryi
(Mollendorff) (fig. 89a-c). Russatus nigrescens (Moll-
endorff) has fewer whorls, a looser coiling pattern, and
is much larger in size (fig. 89d-f). Differences from the
Kusaie Island Sinployea kusaieana (fig. 63a-c) are
covered in the generic discussion above.
Description. Shell rather small, with slightly more than 4'A
relatively tightly coiled whorls. Apex and spire flat, last whorl barely
SYSTEMATIC REVIEW
TABLE XLV. - RANGE OF VARIATION IN MICRONESIAN CHAROPINAE, I.
209
NUMBER OF
NAME
SPECIMENS
RIBS
RIBS/MM
HEIGHT
DIAMETER
Slnployea
kusaieana
61
91.1(84-103)
11.25(10.27-13.12)
1.32(1.19-1.49)
2.60(2
.52-2
.75)
Trukcharopa
trukana
405
150.1(112-197)
17.7(14.07-20.84)
1.19(0.99-1.42)
2.66(2
.50-3,
.08)
Kubaryiellus
kubaryi
52
112.2(99-132)
9.54(8.39-11.70)
1.86(1.59-2.25)
3.74(3
.36-4
.60)
Russatus
nigrescens
30
R E
D U C E D
2.20(1.91-2.66)
4.81(4
.34-5.
53)
Roimontis
tolotomensis
2
57.5(57-58)
6.64(6.27-7.01)
1.46(1.39-1.52)
2.78(2
.65-2,
.91)
Palikirus
cosmetus
8
18.9(17-21)
2.51(2.29-2.71)
1.23(1.12-1.38)
2.40(2
.17-2.
58)
ponapicus
1
46
6.74
0.99
2.17
H/D RATIO WHORLS
kusel. 0.506(0.462-0.542) 3 7/8+(3 3/4-4 1/8)
truka. 0.450(0.371-0.513) 4 3/8-(4 1/8-4 3/4)
kubar. 0.494(0.453-0.540)
nigres. 0.455(0.418-0.482)
tolot. 0.524(0.523-0.525)
cosme . 0.510(0.487-0.542)
ponap. 0.454
4 l/8-(3 7/8-4 1/2)
3 l/8+(3 1/8-3 1/2)
4
3 3/4+(3 5/8-4)
3 3/8
UMBILICUS D/U RATIO
0.74(0.69-0.79) 3.51(3.21-3.67)
0.77(0.63-0.95) 3.43(3.07-4.00)
1.01(0.83-1.32) 3.70(3.36-4.24)
0.53(0.30-0.69) 10.6(6.30-16.5)
0.71(0.69-0.72) 3.91(3.81-4.00)
0.70(0.63-0.82) 3.44(3.14-3.67)
0.54 4.00
APICAL
CORDS
SPIRE
ELEVATION
BODY WHORL
WIDTH SP/BWW
PR
kusei.
9.2(8-10)
0.119(0.099-0.164)
0,
72(0,
79-0
.86) 0.150(0.125-0.208)
truka.
kubar.
9.68(8-12)
7.62(6-9)
REDUCED
6.50(6-7)
11.80(10-13)
9
0.08(FLAT-0.23)
DEPRESSED
SLIGHTLY DEPRESSED
TO FLAT
DEEPLY DEPRESSED
0.17(0.15-0.20)
NOT
0,
1,
1,
1.
A
80(0,
41(1.
29(1
13(1.
.76(0
V A
72-0
,32-1
.18-1
07-1
.72-0
I
.87) 0.092(FLAT-0.264)
co\
2
1
41 1
nigres.
tolot.
-i o \
. 18,1
.79) 0.228(0.205-0.250)
L A B L E
cosme .
ponap .
descending, H/D ratio 0.430. Embryonic whorls l'/2, sculpture of 8
prominent, rather closely spaced spiral cords. Postnuclear whorls
with prominent, narrow, crowded, protractively sinuated radial ribs,
133 on the body whorl, whose interstices are 2-3 times their width.
Microsculpture of very fine radial riblets and even finer spiral rib-
lets, crossed by prominent, widely spaced secondary spiral cords that
become wider and more numerous on the body whorl. Sutures deep,
whorls evenly rounded above, markedly flattened laterally above
periphery and on basal margin. Umbilicus broadly U-shaped, regu-
larly decoiling, contained 3.59 times in the diameter. Color reddish
brown. Aperture ovate, flattened basally and laterally above
periphery, inclined about 5 from the shell axis. Height of holotype
1.12 mm., diameter 2.60 mm.
Holotype. Caroline Islands, Truk, Tol Islet, Sta-
tion A5, on dead leaves, 1,200 m. inland, near top of
Mt. Urupot. Collected by K. Atoda on February 17,
1940. BPBM 189318.
Range. Truk Group and Lukunor, Caroline Is-
lands.
Paratypes. Truk: Moen Islet: Nebokos (Station
16), from shore to 1 A mile inland at 4-5 ft. elevation (4
specimens, BPBM 153701-2); west side of Mt. Teroken
(Station 18), 1 mile inland at 900 ft. elevation (2 speci-
mens, BPBM 153792); Saboku Village (Station A4),
under dead leaves (105 specimens, BPBM 189263).
Fefan Islet: Urunna village (Station A3), near shore on
dead leaves (26 specimens, BPBM 189179). Dublon
Islet: Mt. Tolowan (Station 9), in native forest at 850 ft.
elevation (53 specimens, BPBM 153385-92); Mt. Tolo-
wan (Stations 1, 8, 11) at 300-800 ft. elevation (3
specimens, BPBM 153072, BPBM 153451); Meseran
Village (Station 12), northeast part of island, 1-100 ft.
inland at 1-12 ft. elevation (5 specimens, BPBM
153506). Param Islet: 200 ft. back of beach (Station 37)
at 3 ft. elevation (1 specimen, BPBM 155749). Tarik
Islet (Stations 38, A2; 3 specimens, BPBM 153911,
210
SOLEM: ENDODONTOID LAND SNAILS
BPBM 155762, BPBM 189423). Tol Islet: Mt. Urupot
(Stations 28, 29, 30, A5, A7), halfway up to summit,
1,000-1,200 m. inland, 700-1,422 ft. elevation (109
specimens, BPBM 155550-4, BPBM 153879-82,
BPBM 155477-8, BPBM 155508, BPBM 189318,
BPBM 189343); Urifei (Station A6), on dead leaves (9
specimens, BPBM 189334); Iruku, Tsukuran village
(Station Al), 100 m. inland on dead leaves (13 speci-
mens, BPBM 189406); southwest slope of Mt. Urupot, 1
mile inland above spring, 1,000-2,000 ft. elevation (35
specimens, BPBM 210864, BPBM 210890-2, BPBM
210921). Oneap Islet (9 specimens, ANSP 191994).
Nama Islet (2 specimens, ANSP 191995). Ta Islet (25
specimens, ANSP 192000). Lukunor (3 specimens,
AMSC.7561).
Remarks. Probably Trukcharopa trukana occurs
on most of the islets in the Truk Group. Numerous
specimens were collected on Moen, Fefan, Dublon, and
Tol Islets. Only three living specimens were found on
Tarik and one dead shell from Param Islet. Several sets
from the Thaanum collection were obtained on ex-
change by the ANSP. These sets undoubtedly contain
selected fragments of larger field samples, and the
slight size and shape differences shown by these (table
XLVI) probably reflect sample bias rather than any
morphologic changes. The cited names of Oneap,
Nama, and Ta Islets are taken from the ANSP labels. I
was not able to identify the position of Oneap and Ta.
Ignoring the differences in the ANSP samples,
there is only minor shape variation present. Compar-
ing the specimens from Tol and Moen (table XLVI), the
differences in height (t = 3.6106 with 21 degrees of
freedom), H/D ratio (t = 2.8764), and D/U ratio (t =
2.3826) reach statistical significance. The shells from
Tol all have a flat spire and hence a lower H/D ratio,
lower height, and slightly wider umbilicus than those
from other islets where some individuals have a
slightly elevated spire. I attach no systematic impor-
tance to this variation. Specimens from Tol have much
fewer and more widely spaced major radial ribs than do
those from Moen or Oneap. Shells from Ta (ANSP
192000) are intermediate. Possibly there is an ecologi-
cal correlation involved, but I have no data concerning
this situation.
Where large numbers of specimens were taken,
the percentage of adults is very low. Probably the
greater number of adults in small samples reflects col-
lecting bias, at least in part, but the pattern of juvenile
predominance is clear. Data for six stations are pre-
sented in Table XLVII. The collecting dates range from
December to May and from 1935 to 1940. Two different
collectors were involved. No pattern of seasonality is
suggested.
Four specimens from Lukunor (AMS) are referred
here with some hesitation. They appear to be slightly
subadult, and if allowance is made for this, they fit
within the range of variation for Trukcharopa trukana.
Pending receipt of more adequate material, I classify
these shells as this species.
Several penes were opened, but I am not certain of
the details shown in Figure 90j concerning the exact
stimulatory pad structure. All specimens had many
bits and strands of material inside, and determination
of details was not possible from gross dissections. Time
did not permit preparation of serial sections. The
difference between the Sinployea-type stimulator and
that of Trukcharopa trukana, the former a pocket be-
hind and above a U-shaped muscle band, with the lat-
ter a longitudinally oriented structure, combined with
the presence of an epiphallus in Sinployea and its ab-
sence in Trukcharopa are sufficient to distinguish the
two genera without difficulty.
Description of soft parts. Foot and tail about % shell diameter
in length, not tapering posteriorly, broadly rounded behind, trun-
cated anteriorly. Sole undivided, pedal grooves high on foot, pedal
much deeper, grooves uniting over tail with slight depressions, but
no caudal horn or middorsal groove developed. Slime network with-
out unusual features. Gonopore in normal position. Body color light
yellow-white, without darker markings.
Mantle collar without glandular extension onto pallial roof.
Anus and external ureteric pore with same position. Pallial region
extending V4 whorl apically. Lung roof clear. Kidney about 1.2-1.3
mm. long, rectal arm longer than pericardial arm, which is about 1
mm. from anterior end to base. Ureter typical, section between kid-
ney arms tightly compacted, overlapped by kidney. Heart about half
length of pericardial kidney arm. Principal pulmonary vein un-
branched. Hindgut typical.
Ovotestis (fig. 90i, G) with two clumps of palmately clavate al-
veoli, located in half whorl above stomach apex, lying parallel to
whorl margins and occupying most of whorl space, slightly iridescent
in tone. Hermaphroditic duct partly convoluted, tapering apically,
constricted just before entering base of talon head. Albumen gland
(GG) short and compact, acini relatively small. Talon (GT) with
globular head, very short shaft to carrefour region. Prostate (DG) of
comparatively few and large acini opening into groove on inner side
of uterus. Uterus (UT) bipartite, lower chamber much larger and
with thick, glandular walls.
Vas deferens (VD) slender for entire length, passing through
penial retractor muscle while crossing head of penis, then inserting
laterally on penis head. Penial retractor (PR) arising from dia-
phragm, slightly shorter than expanded penis head, inserting on
head of penis after enfolding vas deferens. Penis (P) with expanded
globular head, about 1.3 mm. long, followed by a narrow tube that is
equal in length to penis head or slightly shorter, internally (fig. 90j)
expanded head with an almost hemispherical stimulator (PP) lying
partly unattached against one wall of penis, no vergic papilla, vas
entering through a simple pore. Internal structure quite complex and
should be studied using serial sections. Atrium (Y) narrow and
rather long.
Free oviduct (UV) larger than penis head, wider above. Sper-
matheca (S) with elongately oval head, very slender shaft, strongly
expanded laterally. Vagina (V) usually longer than illustrated, a
simple tube. Free muscle and digestive system without unusual fea-
tures.
(Based on BPBM 189263, 2 whole individuals 2.43 mm. in
diameter with 4Vt whorls and 2.57 mm. in diameter with 4V4 +
whorls, plus several extracted fragments.)
Genus Kubaryiellus, new genus
Shell large, with about 4'/s rather loosely coiled whorls. Apex
and spire depressed, body whorl not descending. Apical sculpture of
6-9 very prominent spiral cords. Postnuclear whorls with narrow,
prominent, protractively sinuated radial ribs, averaging less than
10/mm. on the body whorl. Microsculpture of prominent radial rib-
lets, finer spirals, and prominent secondary spiral cords. Umbilicus
moderately open, V-shaped, margins rounded. Whorls flattened lat-
SYSTEMATIC REVIEW
211
NAME
NUMBER OF
SPECIMENS
TABLE XLVI. - LOCAL VARIATION IN TRUKCHAROPA.
RIBS /MM.
H/D RATIO
.
Oneap
ANSP 191994
Nama
ANSP 191995
Ta
ANSP 192000
Dub Ion Sta. 9
BPBM 153385, -506
Tol Sta.A5,A6,A7,30
BPBM 155550, BPBM
189318.-34.-43
Moen Sta.A4, 16
BPBM 153701,
BPBM 189263
Fefan Sta. A3
BPBM 189179
1
Lukunor
AMS C7561
11
11
Oneap
Nama
Ta
Dub Ion
Tol
Moen
Fefan
Lukun .
168.817.46
(143-197)
19.0+0.550
(16.98-20.50)
1.2710.032
(1.19-1.39)
2.8510.055
(2.70-3.08)
0.45210.0034
(0.442-0.460)
157
17.98
1.1310.017
(1.12-1.15)
2.7010.083
(2.62-2.78)
0.42110.0065
(0.414-0.427)
139.313.14
(124-156)
17.310.359
(15.07-19.24)
1.21+0.019
(1.08-1.27)
1 . 16+0 .017
2.5710.023
(2.45-2.68)
2 64+0 029
0.46910.0078
(0.420-0.506)
440*0 0091
(1.09-1.23)
(2.52-2.75)
(0.398-0.468)
122.5
(112-133)
15.2
(14.09-16.28)
1.1410.024
(0.99-1.29)
2.64+0.036
(2.52-2.95)
0.433+0.0090
(0.371-0.487)
164.7
(147-183)
18.5
(16.54-20.80)
1.2610.023
(1.16-1.42)
2.7110.041
(2.55-2.91)
0.466+0.0069
(0.432-0.513)
(1.19-1.32)
(2.58-2.88)
(0.437-0.471)
114.8+2.78
(107-120)
16.610.859
(15.01-19.02)
1.0610.032
(0.97-1.12)
2.20+0.067
(2.01-2.30)
0.480+0.0052
(0.471-0.493)
WHORLS
UMBILICUS
D/U RATIO
APICAL
CORDS
4 l/2-(4 1/4-4
3/4) 0.86
(0.76-0,
3.2910.074
,95) (3.07-3.59)
10.67+0.335
(10-12)
4 1/4
0.81+0.050
(0.76-0.
3.3610.105
86) (3.25-3.46)
10.0
4 l/4-(4 1/8-4
3/8) 0.7510.015
(0.63-0,
3.4210.066
82) (3.18-3.97)
10.36+0.244
(9-12)
4 l/4+(4 1/8-4
1/2) 0.7410.023
(0.63-0,
3.5710.087
79) (3.29-4.00)
9.8610.261
(9-11)
4 3/8-(4 1/4-4
5/8) 0.7910.019
(0.72-0,
3.33+0.052
95) (3.07-3.63)
8.7310.305
(8-11)
4 3/8-(4 1/8-4
5/8) 0.7810.017
(0.69-0.
3.4810.033
86) (3.27-3.67)
9.2010.250
(8-10)
4 l/2-(4 1/4-4
1/2) 0.76+0.028
(0.69-0.
3.5710.058
82) (3.40-3.71)
9.7510.480
(9-11)
4-(3 3/4-4 1/8)
0.5810.028
(0.53-0.
3.79+0.140
66) (3.40-4.06)
erally above and below rounded periphery, aperture inclined about
15 from shell axis. No apertural barriers. Pallial cavity with rectal
kidney arm about twice length of pericardial, otherwise without un-
usual features. Ovotestis with one clump of alveoli, hermaphroditic
duct tightly convoluted. Vas deferens passing through penial retrac-
tor muscle before entering laterally on penis head. Penis very long,
club-shaped, tapering, internally with a trilobed vergic papilla fol-
lowed by 2 long, stimulatory pilasters on a papillose glandular wall.
A thin muscle sheath surrounds penis. Spermatheca typical, vagina
somewhat longer than typical.
TABLE XLVII. - AGE DISTRIBUTION IN TRUKCHAROPA TRUKANA.
ISLET
Moen
Dub Ion
Tol
STATION
DATE OF NUMBER
COLLECTION OF ADULTS
NUMBER
OF SHELLS
COLLECTED
% OF
ADULTS
A3
A4
May
Mav
1, 1940 5
24, 1940 9
26
105
19
8
'
6%
9
Dec.
20. 1935
53
13
.2*
A5
A6
30
Feb.
Mav
17, 1940 5
9, 1940 2
4, 1936 3
24
9
47
20
22.
6
-
2%
.
Type species. Charopa kubaryi Mollendorff,
1900.
Roimontis tolotomensis (fig. 91a-c) is most similar
in shell features, but differs in having spiral grooves
rather than spiral cords, two parietal barriers, fewer
and more widely spaced radial ribs, a much more de-
pressed spire, and it is smaller in size. Russatus nigres-
cens (fig. 89d-f) has a flat spire, much looser coiling, a
very narrow umbilicus, reduced radial sculpture,
greatly altered pallial region, and quite different geni-
tal structures. Trukcharopa trukana (fig. 63d-f) has a
flat to slightly elevated spire, much more numerous
and crowded radial ribs on a much smaller shell, a
tighter coiling pattern, finer apical sculpture, and the
penis not only lacks a muscle sheath but has much
different internal structure.
Kubaryiellus (fig. 89a-c) has a modified shell. Its
depressed apex and spire with relatively loose coiling
are unusual in the Pacific Island Charopidae, but the
212
SOLEM: ENDODONTOID LAND SNAILS
systematically important features are the greatly al-
tered penial complex. This agrees with most other Mi-
cronesian genera in having the vas deferens pass
through the penial retractor muscle, but the distinct
penial muscle sheath, trilobed vergic papilla, and two
enlarged longitudinal pilasters (fig. 90g) represent
major departures from the structural pattern seen in
the other genera. The penial length of 1.95-2.20 mm. is
much greater than that seen in the other Micronesian
genera that lack an epiphallus. The vergic papilla
could be derived from the type seen in Sinployea ir-
regularis (fig. 67d) by elongation and size increase, but
the absence of an epiphallus, development of a distinct
sheath, and the peculiar pilasters are very different
from the Sinployea pattern.
Kubaryiellus is named in honor of I. Kubary,
pioneer collector of land mollusks in New Guinea and
the Caroline Islands for Otto von Mollendorff.
Kubaryiellus kubaryi (Mollendorff, 1900). Figures
89a-c, 90e-h.
Charopa kubaryi Mollendorff, 1900, J. Malacol., 7 (5), p. 110
Ponape, Caroline Islands.
Diagnosis. Shell large, diameter 3.36-4.60 mm. (mean 3.74
mm.) with 3 7 /-4'/2 normally coiled whorls. Apex and spire distinctly
depressed below level of body whorl, last portion of body whorl barely
descending, H/D ratio 0.453-0.540 (mean 0.494). Apical sculpture of
6-9 (mean 7.62) prominent spiral cords, whose interstices are about
twice their width. Postnuclear sculpture of prominent, narrow,
lamellar, crowded, protractively sinuated radial ribs, 99-132 (mean
112.2) on the body whorl, whose interstices are 3-6 times their
width. Ribs/mm. 8.39-11.70 (mean 9.54). Microsculpture of promi-
nent radial riblets, 3-6 between each pair of major ribs, much finer
and more crowded spiral riblets, with rather widely spaced secondary
spiral cords equal in size to or slightly larger than microradial rib-
lets. Umbilicus broadly V-shaped, regularly and rapidly decoiling,
contained 3.36-4.24 times (mean 3.70) in the diameter, margins
evenly and gently rounded. Sutures deep, whorls strongly rounded
above and on baso-columellar margin, strongly compressed laterally
above rounded periphery with slightly compressed lower palatal
margin. Aperture subovate, strongly compressed laterally above
periphery, inclined about 15 from shell axis. No callus or apertural
barriers developed.
In having a distinctly depressed apex and marked
lateral flattening of the body whorl, Kubaryiellus
kubaryi is immediately separated from other Microne-
sian or Polynesian species. Roimontis tolotomensis (fig.
91a-c) has the apex much more deeply depressed, two
weak parietals, and is much smaller.
Description. Shell large, with 4V4 overlapping whorls. Apex
and early spire distinctly sunken, markedly overlapped by coiling of
later whorls, last portion of body whorl descending only slightly
below top of penultimate, H/D ratio 0.458. Apical whorls 1H,
sculpture of 7 widely spaced, relatively narrow spiral cords. Postnu-
clear whorls with somewhat irregular, narrow, moderately protrac-
tively sinuated radial ribs, relatively widely spaced on spire, becom-
ing crowded on last whorl, about 132 on body whorl, whose
interstices are 1-3 times their width. Microsculpture of fine, irregu-
larly spaced radial riblets crossed by finer, slightly more crowded
spiral riblets with a secondary sculpture of moderately prominent,
widely spaced spiral cords. Sutures moderately impressed, whorls
strongly rounded above, strongly flattened laterally above obtusely
rounded periphery with evenly rounded basal margin. Umbilicus
broadly V-shaped, regularly decoiling, contained 3.43 times in the
diameter, with margins slightly shouldered. Color light reddish
yellow-brown above, apex yellow-horn, and a narrow yellowish white
spiral band situated Vs of the way between suture and periphery of
the whorl. Periostracal ends of ribs in sutures and in umbilicus dark
chocolate brown. Aperture ovate, moderately flattened laterally
above periphery, inclined about 10 from the shell axis. Height of
lectotype 1.81 mm., diameter 3.95 mm.
Lectotype. Caroline Islands: Ponape. Collected by
Kubary. SMF 165352.
Range. Ponape, Caroline Islands.
Paratypes. BPBM 86245, SMF 165353.
Mater ial Ponape (1 specimen, BPBM 87528):
back of Naupilo (Station 133) (4 specimens, BPBM
157564-5); Paishapel Hill, Metalanim (Station 113), at
650-700 ft. elevation (14 specimens, BPBM 157211-6);
west side Tolomaim Mt. (Station 91) at 1,000-1,500 ft.
elevation (3 specimens, BPBM 154002-3); Metalanim
to Mipit (Station 118), 450 ft. inland at 300-900 ft.
elevation (6 specimens, BPBM 154157-60); Mt.
Kuporujo (Station 121) at 500-900 ft. elevation (1
specimen, BPBM 154338); southwest side of To-
reairuka Mt. (Station 127) at 250-750 ft. elevation (6
specimens, BPBM 154451-3); one quarter to halfway
up Mt. Tolotom (Station 92) at 800 ft. elevation (1
specimen, BPBM 156677); north slope Mt. Nanalaut
(Stations 129, 130) at 1,600-2,500 ft. elevation (2
specimens, BPBM 157443); Mt. Nanalaut (Station
129A) at 1,900 ft. elevation to top (1 specimen, BPBM
154523); Palikir, Jokaj (Station 139 = 138), at 600-900
ft. elevation (3 specimens, BPBM 157797); Palikir, Mt.
Tamantamansakir (Station 140) at 1,000-1,250 ft. ele-
vation (6 specimens, BPBM 157864).
Remarks. Relatively few individuals of Ku-
baryiellus kubaryi were taken at any single station,
but it is widely distributed on Ponape. From stations
where individuals were measured, 39 shells were col-
lected, of which 20 (51.3%) were adult. This is by far
the highest percentage of adult specimens for any
Caroline species of which more than 10 examples were
collected.
Spire depression is marked, but much less than in
Roimontis tolotomensis, and there is no noticeable re-
duction in umbilical depth.
Dissection of several penes showed variation in the
length and exact form of the vergic papilla. The figured
example (fig. 90g, PV) had the shortest seen, the lower
lobes are sometimes extended for almost half the penis
length in a U-shaped form, the upper lobe being rela-
tively constant in shape. In all examples the opening of
the vas deferens was beneath the median lobe of the
vergic papilla, the two lower and lateral lobes serving
as a channel for sperm transfer.
No systematic significance should be given to the
great size variation shown by the individual small sets
measured (table XLVIII). Because of the depressed
spire, already crowded ribbing, regular umbilical de-
coiling, and lack of body whorl descension, separation
of adult from subadult shells is virtually impossible.
Probably the specimens from Stations 113 and 127 are
fairly close to typical adult size. The types are dis-
def
a be
FIG. 89. a-c, Kubaryiellus kubaryi (MbllendorfD. Ponape, Caroline Islands. Paratype. BPBM 87528 ex SMF; d-f, Russatus nigrescens
(MollendorfT). Station 129A, Mt. Nanalaut, Ponape, Caroline Islands. BPBM 154521. Scale lines equal 1 mm. (MM).
213
214
SOLEM: ENDODONTOID LAND SNAILS
tinctly smaller and more widely umbilicated than the
Micronesian Expedition shells. This is not a size-
correlated change, since almost certainly subadult
shells from Station 140 agree in umbilical width with
the other Micronesian Expedition shells. The latter
were collected at higher elevations than the types
(Mollendorff, 1900, p. 101). The opposite variation is
seen in Russatus nigrescens (table XLVIII), where the
low elevation syntype specimens were much more nar-
rowly umbilicated than the Micronesian Expedition
upland shells. Further collections of both species are
needed.
Description of soft parts. Foot and tail long and slender, up to
3.75 mm. long, tapering posteriorly, broadly rounded at end, trun-
cated in front. Sole undivided. Pedal grooves high on foot, pedal
much stronger, both uniting above tail, but no caudal horn or mid-
dorsal groove present. Slime network details not clear in available
material. Head partly retracted in all specimens examined. Gono-
pore position not observed. Body color light yellow-white, without
darker markings.
Mantle collar without glandular extension onto pallial roof.
Anus (A) opening at inner edge of pneumostome, external ureteric
pore (KX) opening well behind anus. Pallial region (fig. 90e) about
3.6 mm. from inner edge of mantle collar to apex of kidney. Lung roof
clear, without granulations. Kidney (K) with 2 lobes, pericardial
barely longer than heart, rectal arm about 2.2 mm. long, more than
twice length of pericardial arm. Ureter (KD) with arms closely ap-
proximated, no lung roof visible between, tapering anteriorly. Heart
(H) lying parallel to hindgut, about equal in length to pericardial
kidney lobe. Principal pulmonary vein (HV) without obvious branch-
ing. Intestinal loop just above pallial cavity apex, hindgut (HG)
reaching parietal-palatal margin just above pallial cavity apex.
Ovotestis (fig. 90f, G) one clump of 5-8 iridescent, very elon-
gated acini, lying parallel to sides of whorls, located just above
stomach apex, extending a fraction of a whorl. Hermaphroditic duct
(GD) tightly convoluted apically and, in some specimens, just before
talon, iridescent for most of length, inserting at base of talon head.
Albumen gland (GG) short, compact. Talon (GT) partly buried in
surface of albumen gland, head globose, shaft slender to carrefour.
Prostate (DG) of narrow acini opening into groove on inner surface of
uterus, acini variable in length. Uterus (UT) bipartite, upper
chamber much more thin-walled and more slender than thick-walled
lower chamber (fig. 90h).
Vas deferens (VD) thin-walled at first, becoming more muscular
at penioviducal angle, loosely or not bound to side of penis, entering
penial apex after passing through penial retractor muscle and
traversing head of penis. Penial retractor (PR) very short, arising
from diaphragm, inserting on head of penis after enfolding vas defer-
ens. Penis (P) about 1.95-2.20 mm. long, club-shaped, covered by a
thin muscle sheath, usually tapered basally (figured example
bifolded basally), internally (fig. 90g) with a trilobed vergic papilla
(PV) through which vas deferens enters. Lower part of penis appar-
ently with two simple to corrugated longitudinal pilasters that form
one side of the papillate lower penial region. Atrium (Y) long and
much more slender than penis base.
Free oviduct (UV) shorter than penis, wider apically and slightly
expanded before joining spermathecal shaft. Spermatheca (S) with
ovate head next to albumen gland, shaft slender to level of free
oviduct, then greatly expanded with thick glandular walls. Vagina
(V) elongated, slender, with weak internal pilasters. Free muscle
system without unusual features. Salivary glands fused posteriorly
above esophagus.
(Based on BPBM 157211-2, 5 extracted and partial specimens.)
Genus Russatus, new genus
Shell very large, with less than 3'/2 very loosely coiled whorls.
Apex and spire flat or slightly depressed, body whorl descending
slightly. Apex macroscopically smooth, traces of fine spiral cording
visible under high magnification. Postnuclear whorls with irregular,
very crowded, fine radial ribs with high periostracal extensions, on
lower spire not separable from growth irregularities. Occasional
microreticulation visible at 96 x magnification. Umbilicus very nar-
row, V-shaped, regularly decoiling. Whorls compressed laterally
above periphery and on basal margin, aperture strongly inclined
from shell axis. No apertural barriers. Pallial cavity extending 1 A
whorl apically, kidney almost square, very thick in middle, rectal
and pericardial arms equal in length. Apical genitalia not examined.
Penial retractor muscle enfolding vas deferens before latter enters
laterally on penis head through a simple pore. Expanded head of
penis triangular, internally with a blind caecum lying at apex from
opening of vas deferens to outer edge of expanded penis head, lower
parts with circularly elongated glandular papillae on the walls.
Spermatheca with basal part of stalk globularly expanded, vagina
relatively long and much thicker than free oviduct.
Type species. Flammulina (Calymna) nigrescens
Mollendorff, 1900.
Except for the Guam Island Himeroconcha fusca
(fig. 105d-f), Russatus nigrescens is the largest Mi-
cronesian charopid. Its major distinguishing features
are the reduced whorl count (table XLV), much looser
coiling pattern (fig. 89d), great sculptural reduction,
development of a square kidney (fig. 90a) that remains
next to the hindgut, and the very peculiar caecum on
the inner wall of the penis (fig. 90d). No Polynesian or
Micronesian species is at all similar in shell coiling
pattern, and the whorl count is lower than all known
species. Only the minute and very tightly coiled Mi-
crocharopa mimula (fig. 33a-c) and the very high-
spired Ba humbugi (fig. 74a-c) from Viti Levu, Fiji,
come close in terms of whorl count. They differ mark-
edly in coiling pattern, apical sculpture (Microcharopa,
fig. 4a-d), and genital anatomy (Ba, fig. 75a-h).
The New Guinea endemic Paryphantopsis (Solem,
1970a, fig. la-c, shell, and fig, 2a-e, anatomy) is simi-
lar in whorl count, coiling pattern, major sculpture re-
duction, and size, but the similarities are convergent.
In Paryphantopsis the pallial cavity has the kidney
partly rotated from the hindgut, the arms of the ureter
are separated at a 60 angle instead of being in contact,
the principal pulmonary vein is noticeably branched,
and the entire pallial cavity only occupies one-sixth
whorl. Paryphantopsis has quite different terminal
genitalia, there being an enlarged epiphallus with a
prominent caecum, the penis with a verge and corru-
gated longitudinal pilasters, and the vaginal region
being very short and swollen with thick glandular
walls. Russatus lacks both an epiphallus and a verge.
Species of Paryphantopsis have the umbilicus com-
pletely closed or at most a weak lateral crack, whereas
the umbilicus of Russatus nigrescens is distinctly open.
Both genera have started the process of visceral
hump reduction that can eventually lead to evolution
of a "slug" genus, but the pallial cavity and genital
structures indicate that they are not closely related.
Either Kubaryiellus kubaryi (fig. 90e-h) or Truk-
charopa trukana (fig. 90i-j) might be compared with
Russatus nigrescens as a possible ancestral type. The
first has a large vergic papilla, very prominent
stimulators, and the pallial cavity with very long rec-
tal and much shorter pericardial kidney arms. Truk-
SYSTEMATIC REVIEW
215
TABLE XLVIII. - LOCAL VARIATION IN KUBARYIELLUS AND RUSSATUS .
Kubaryiellus
kubaryl
BPBM 86245,
SMF 165352-3
Sta.113
BPBM 157211-2
Sta. 127
BPBM 154451-2
Sta. 140
BPBM 157864
Russatus
nigrescens
Pre-1900
SMF 165551-2,
DMW MF8753
In 1936
BPBM 154077, -403,
-520, BPBM 156678
117.77.85
(105-132)
132.018.27
(108-168)
RIBS/MM.
9.900.431
(9.15-10.64)
10.13*0.784
(9.34-11.70)
1.78+0.016
(1.74-1.81)
2.00+0.064
(1.88-2.14)
2.03+0.076
(1.88-2.24)
1.71+0.040
(1.58-1.84)
2.150.033
(2.01-2.27)
2.26+0.177
(1.92-2.68)
H/D RATIO
3.73+0.079 0.479+0.0094
(3.59-3.95) (0.458-0.495)
4.09+0.183 0.490+0.0159
(3.68-4.57) (0.453-0.524)
3.97+0.091 0.51110.0102
(3.71-4.14) (0.492-0.540)
3.47+0.058 0.4930.0068
(3.36-3.72) (0.471-0.519)
4.77+0.053 0.450+0.0072
(4.57-5.00) (0.418-0.475)
4.82+0.283 0.4640.0125
(4.37-5.56) (0.430-0.483)
APICAL
D/U RATIO CORDS
4 1/8(4-4 1/4)
Sta. 113
Sta. 127
nigrescens
Pre-1900
In 1936
1.08+0.028 3.46+0.062
(1.02-1.15) (3.36-3.64)
4 3/8- (4 1/8-4 1/2) 1.11+0.072 3.70+0.136 7.50.500
(0.99-1.32) (3.48-4.06) (6-8)
4 l/4-(4-4 1/2)
4+(3 7/8-4 1/8)
3+(3-3 1/8)
1.05+0.036 3.7910.147 6.8+0.4,79
(0.95-1.12) (3.53-4.21) (6-8)
0.9410.041 3.7310.108 8.210.307
(0.82-1.09) (3.43-4.12) (7-9)
0.35+0.040 14.22+1.38
(0.30-0.43) (11.70-16.45)
3 l/4+(3 1/8-3 1/2) 0.62+0.054 7.92+0.685 REDUCED
(0.46-0.69) (6.30-9.64)
charopa has the penis without a vergic papilla, pos-
sesses a very unusual medial penial stimulator and
agrees in pallial configurations with Kubaryiellus. Al-
though their shell form appears intermediate between
the more typical Trukcharopinae and Russatus, their
anatomical features indicate no close affinity.
The name Russatus, meaning clothed in red, refers
to the very dark color of the shell.
Russatus nigrescens (Mollendorff, 1900). Figures
89d-f, 90a-d.
Flammulina (Calymna) nigrescens Mollendorff, 1900, J. Malacol.,
7 (5), pp. 107-108, figs. 1-3 Naupilo. Ponape, Caroline Islands,
at 100 m. elevation.
Diagnosis. Shell very large, diameter 4.34-5.53 mm. (mean
4.81 mm.), with S'/s-S'/a loosely coiled whorls. Apex and spire flat or
very slightly depressed, last part of body whorl descending slightly,
H/D ratio 0.418-0.482 (mean 0.455). Apical whorls with faint traces
of spiral cords under high magnification, macroscopically smooth.
Postnuclear whorls with irregular, extremely narrow and crowded,
protractively sinuated radial ribs with lamellar extensions, becom-
ing indistinguishable from growth irregularities shortly after apex.
Occasional traces of microreticulation visible under 96 x magnifica-
tion. Umbilicus very narrow, broadly V-shaped, regularly decoiling,
contained 6.30-16.5 times (mean 10.6) in the diameter, margins
strongly rounded. Sutures deep, whorls strongly rounded above,
greatly compressed laterally above strongly rounded periphery and
on basal margin. Aperture elongately ovate, compressed laterally
above periphery and on basal margin, inclined almost 40 from shell
axis. No apertural barriers.
Russatus nigrescens has the lowest whorl count
and loosest coiling pattern of all Pacific Island En-
dodontidae and Charopidae. Its large size, tiny um-
bilicus, reduced apical sculpture, and very crowded,
lamellar major radial sculpture immediately separate
it from any Micronesian species.
Description. Shell very large, with 3!/s very loosely coiled
whorls. Apex and early spire flat, body whorl descending very
slightly, H/D ratio 0.466. Apical whorls IVfe, sculpture eroded with
only faint traces of spiral cording visible in sutures. Postnuclear
whorls with irregularly spaced radial ribs and growth threads plus a
fine wavy spiral sculpture of riblets. Sutures moderately impressed,
whorls flatly rounded above and on basal margin. Color dark
purplish brown with occasional lighter patches becoming deep purple
in the sutures and umbilicus. Umbilicus very narrow, greatly con-
stricted by growth of periostracum, contained 16.45 times in the
diameter. Aperture elongately ovate, flattened above periphery and
on basal margin, inclined almost 35 from the shell axis. Height of
lectotype 2.27 mm., diameter 4.87 mm.
216
SOLEM: ENDODONTOID LAND SNAILS
PP
FIG. 90. Anatomy of Russatus, Kubaryiellus, and Trukcharopa: a-d, Russatus nigrescens (Mollendorff). Station 129A, Ponape. BPBM
154522. a, pallia! region, b, genitalia. c, interior of penis with caecum (PC) hiding vas opening, d, interior of penis showing relation of vas
opening (DP) and pore of caecum; e-h, Kubaryiellus kubaryi (Mollendorff). Station 113, Ponape. BPBM 157212. e, pallial region, f, genitalia. g,
interior of penis, h, detail of prostate-uterus ending; i-j, Trukcharopa trukana, new species. Station A4, Moen Islet, Truk. BPBM 189263. ;',
genitalia;j, interior of penis. Scale lines equal 1 mm. (MO).
Lectotype. Caroline Islands: Ponape, Naupilo, at
about 100 m. elevation. Collected by Kubary. SMF
165551.
Range. Ponape, Caroline Islands.
Paratypes.SMF 165552.
Material. Ponape (1 specimen, DMW): Mt. To-
lotom (Station 92), from V4-% way up, near 800 ft. ele-
vation (2 specimens, BPBM 156678); Mt. Tolotom (Sta-
tion 93), % way up, to summit at 1,200 ft. elevation on
dead leaves (11 specimens, BPBM 154077-80); Mt.
Nanalaut (Station 129A), from 1,900 ft. to summit, on
ground (8 specimens, BPBM 154520-2); Mt. Kuporujo,
near summit (Station 122) at 1,600-1,970 ft. elevation
(2 specimens, BPBM 154403).
SYSTEMATIC REVIEW
217
Remarks. Only four of 23 specimens collected by
the Micronesian Expedition were adult, and each came
from a different station. Although the original collec-
tion was made at low altitude. 100 m., subsequent finds
have been at higher elevations from widely scattered
localities. The types are much more narrowly umbili-
cated than the more recently collected shells (table
XLVIII), but I am uncertain as to the significance of
this difference. Possibly subspecific differentiation is
involved.
Suter (in Mollendorff, 1900, p. 108, figs. 1-3) illus-
trated the tail of a dried, then softened, individual, part
of the jaw, and a portion of one radular half row. His
classification of this species with Flammulina was a
logical guess but is not supported by subsequent dissec-
tions. Flammulina shows major differences in pallial
and genital anatomy (fig. 29a-j). The conchological
similarities clearly are convergent.
Russatus nigrescens has fewer whorls, mean count
3Vs + , than any Pacific Island endodontoid. The Fijian
Ba humbugi has somewhat similar alteration in form,
but clearly differs in its greatly elevated spire and
anatomical structure (fig. 75a-h). Ba humbugi does
have a mean whorl count of 3%, but the pattern of
coiling (fig. 74a) is very different.
Description of soft parts. Foot and tail about 5.5 mm. long,
tapering gradually from visceral stalk to narrowly rounded tip, trun-
cated anteriorly. Sole undivided. Pedal groove very prominent, su-
prapedal much weaker, both uniting